Type specimens of recent and fossil Testudines and Crocodylia in the collections of the Netherlands Centre for Biodiversity Naturalis, Leiden, the Netherlands

Type specimens of recent and fossil Testudines and Crocodylia in the collections of the Netherlands Centre for Biodiversity

Naturalis, Leiden, the Netherlands

M.S. Hoogmoed, M.E. Gassó Miracle & L.W. van den Hoek Ostende

Hoogmoed, M.S., M.E. Gassó Miracle & L.W. van den Hoek Ostende. Type specimens of recent and fos-sil Testudines and Crocodylia in the collections of the Netherlands Centre for Biodiversity Naturalis, Leiden, the Netherlands.Zool. Med. Leiden 84 (8), 15.x.2010: 159-199, figs 1-13.― ISSN 0024-0672.Marinus S. Hoogmoed, Museu Paraense Emilio Goeldi, Caixa Postal 399, 66017-970 Belém, Pará, Brazil ([email protected]).Eulàlia Gassó Miracle, Netherlands Centre for Biodiversity Naturalis, P.O. Box 9517, 2300 RA Leiden, The Netherlands ([email protected]).Lars W. van den Hoek Ostende, Netherlands Centre for Biodiversity Naturalis, P.O. Box 9517, 2300 RA Leiden, The Netherlands ([email protected]).

Key words: Testudines; Crocodylia; types; Leiden Museum.We report a total of 57 type specimens (31 primary and 26 secondary) belonging to 27 nominal species of chelonians (23) and crocodilians (4) in the collection of the Netherlands Centre for Biodiversity Natu-ralis Leiden, the Netherlands. Fragments of fossil crocodilians have not been included in the specimen count. We provide the original name and publication, type locality, synonyms under which type speci-mens have been cited in the literature, current name, comments on the type material, its history, and its status and provide information on illustrations.

Introduction

The Rijksmuseum van Natuurlijke Historie (RMNH), presently Netherlands Centre for Biodiversity Naturalis, was founded August 9th, 1820 by King Willem I of The Netherlands. At that time three existing collections were joined: ‘s Landskabinet van Natuurlijke Historie (established in 1808 as Cabinet du Roi and renamed in 1810), the ‘Ancien Cabinet’ containing material from Leiden University (of which the collection of the Stadtholder formed part), and the private collection of the first director of the RMNH, C.J. Temminck (Gijzen, 1938; Holthuis, 1995; Fransen et al., 1997). The origin of this material was rather diverse and sometimes difficult to track, but some of the mate-rial dated back to at least 1750. However, of the chelonians and crocodilians treated here, only one dry tortoise shell formed part of the collections present in 1820. The rest was collected after the establishment of the RMNH. Just after founding the RMNH, at the instigation of C.J. Temminck a multidisciplinary commission (‘Natuurkundige Commissie’) for the scientific exploration of the then Dutch East Indies (now Indonesia) was established, and the first members (H. Kuhl and J.C. van Hasselt accompanied by the artist J. Keultjes and the technician G. van Raalten) travelled to Java in 1820 where they arrived in 1821 (Holthuis, 1995), and immediately started their work of collecting and describing natural products. This commission functioned till 1850, when it was dis-solved and for a number of years thereafter, its work was continued by several travelers that continued to provide the RMNH with material. The members of this commission provided the RMNH with an enormous amount of material (preserved specimens,

160 Hoogmoed et al. Types Testudines and Crocodiles Leiden. Zool. Med. Leiden 84 (2010)

drawings of live specimens and field notes), of which many specimens served for the description of new species. Members of the commission that were responsible for the collection of chelonian and crocodilian type specimens (besides many other herpeto-logical specimens) are H. Kuhl & J.C. van Hasselt (on museum labels generally indi-cated as ‘K & vH’), H. Boie & H.C. Macklot, S. Müller, C.A.L.M. Schwaner, L. Horner, E. A. Forsten and P. Diard. Besides the three basal collections present in 1820, this material from the Dutch East Indies collected in the first years of the existence of the RMNH, formed the backbone of the RMNH collections and still is a rich source of new knowledge. Apart from this concentrated effort in the Dutch East Indies, C.J. Temminck (direc-tor of the RMNH from 1820 until 1858) and his successor H. Schlegel (director from 1858 until 1884) maintained close contacts with Dutch diplomats and civil servants in many Dutch outposts in all corners of the world. Thus important material was also obtained e.g. from southern Africa by the members of the ‘Natuurkundige Commissie’ H. Kuhl and J.C. van Hasselt during a stopover on their way to the East Indies and by J.B. von Horstock, from Ghana through H.S. Pel, from Japan through P.F. von Siebold and H. Bürger, from Suriname through H.H. Dieperink (though no type material here discussed), from the U.S.A. through G. Troost, and from the Dutch East Indies through C.F.H. ten Kate. Salomon Müller was one of the two members of the ‘Natuurkundige Commissie’ that returned to the Netherlands and was able to publish part of the scientific results of his stay in the Dutch East Indies, either by himself (Müller, 1838), or together with Schlegel (Müller & Schlegel, 1839; 1841; Schlegel & Müller, 1840, 1845). From Müller’s publications (and other documents about the work of the ‘Natuurkundige Commissie’) it becomes clear what pioneering work was done by the members of the ‘Natuurkun-dige Commissie’, most of which unfortunately could not publish their results (of the 18 persons sent out as members of this commission most died in the East Indies and only two returned to the Netherlands) (Holthuis, 1995). Immediately after the foundation of the RMNH the director C.J. Temminck al-ready maintained good contacts with other major museums in Europe and instigated a lively exchange policy, mainly using material from the Dutch East Indies, Suriname and Ghana to obtain material from parts of the world from which the RMNH only had small or no collections. Museums involved in these exchanges are those from Munich (material collected by J.B. von Spix), Paris (Testudo dussumieri), Berlin, Vienna and London, to name just a few of the most important ones. Thus, part of the type series of chelonians (and other reptiles and amphibians as well) have been split up between several European museums, because the concept of type specimens in the early 19th century was not as well developed as it is now, and series of specimens used for a type description were just considered as ‘doubletten’ (doubles) and not given special status and consequently freely used for exchange. This exchange policy was maintained throughout the history of the herpetological collection, but later was based on modern opinions about the value of material, especially type material. Che-lonian material also reached the museum via other channels: three type specimens (a paratype of Geoemyda spengleri sinensis and two of Batrachemys heliostemma were ob-tained as gifts from foreign researchers, three paratypes (Cyclemys pulchristriata and two specimens of Emys orbicularis occidentalis) were obtained from animal dealers, and

Hoogmoed et al. Types Testudines and Crocodiles Leiden. Zool. Med. Leiden 84 (2010) 161

one paratype (E. orbicularis occidentalis) was collected during fieldwork in 1967 by the senior author. Most of the material of recent and fossil species here reported was collected in the 19th century (most chelonians and all crocodilians), and most of the descriptions also are from that century. One specimen (Testudo indica Vosmaeri) was collected and described in the 18th century, only a few specimens of chelonians were collected in the 20th century. Five descriptions of chelonians were published in the 20th century and two were pub-lished in the 21st century. All fossil material here reported was collected by E. Dubois in the Dutch East Indies near the end of the 19th century and described in the early 20th century.

Material and methods

Part of the material dealt with here is preserved in alcohol and part is preserved as dry material: skeletons (complete or partial [mostly skulls]), skins and stuffed material. Van Lidth de Jeude (1898) reported on part of the dry material (skeletons, skulls and skeletonised specimens of turtles and tortoises with the horny scales still attached to the carapace) present at that time. Stuffed material or skins was not included. This publica-tion in the RMNH is popularly known as the ‘Catalogue Ostéologique’ and material mentioned in it has labels with the indication ‘Cat. ost.’ followed by a letter correspond-ing with the letters mentioned in van Lidth de Jeude (1898). Part of this material in the meantime has been relabelled and now has serial RMNH numbers. The Cat. ost. letters are mentioned after the present serial number. However, the old Cat. ost. indication in all specimens remains in place, although it is superseded by the serial number for easi-er access and incorporation into the electronic database. The original name and publication on which a nominal taxon is based have been indicated by an asterisk preceding the name. References in synonymy lists only concern publications which specifically mention RMNH type material. Thus, no reference is made to the most recent checklist of Fritz & Havaš (2007) under each species, but for present day Chelonian nomenclature we did follow this publication, with the exception of one name (Dipsochelys dussumieri) due to new data that were not taken into account in this publication, although the basic publications concerning the nomenclature of this species were referred to. For crocodile nomenclature we followed King & Burke (1989) and Ross (1990, 1992). Current names for taxa are indicated in bold letters, either in the caption of each account, when the name did not change, or in a special section ‘Current name’. Collection acronyms are based on Frost (2009). The former Rijksmuseum van Na-tuurlijke Historie has gone through several name-changes and at the moment is offi-cially registered as Foundation Nationaal Natuurhistorisch Museum, but the acronym RMNH (which refers to recent material and is recognized internationally) has been maintained in order to avoid confusion. In order to adhere to international standards, the herpetological collection numbers are now indicated as RMNH.RENA. The nota-tion ‘alc.’ means preserved in ethanol 70%, ‘RMNH. Dub.’ refers to the RMNH Dubois collection of fossils. Names of fossil species are preceded by a †. As usual in many old collections, type specimens often were not indicated as such, and only starting in the 1930s the curators of the RMNH herpetological collection


(L.D. Brongersma (1932-1958), M. Boeseman (temporarily supervising the herpeto-logical collection between 1958 and 1966) and M.S. Hoogmoed (1966-2004) [the years mentioned by Gassó et al., 2007 are not correct]) made it a habit of clearly marking type material with red dots on bottles and pedestals, and noting these findings in the card-index and in the hand written register books, whenever they found indications material had been used for descriptions of new taxa. For chelonians and crocodiles we are convinced that all type material now has been identified, and that the present list is complete.

TESTUDINESCRYPTODIRA

Chelydridae Gray, 1831Chelonura Temminckii Troost in Harlan, 1835

*Chelonura Temminckii Troost in Harlan, 1835: 157-158.Macroclemys temminckii; King & Burke, 1989: 26.

Holotype: possibly RMNH.RENA 6166, 1 mounted ex. Loc.: ‘Tennessee’. Leg.: G. Troost. Received in 1839.

Current name: Macrochelys temminckii (Troost in Harlan, 1835). Remarks.― The name Chelonura Temminckii was taken from a manuscript by Troost written in 1835 (Harlan, 1835: 158; Holbrook, 1842: 151). According to Bour (1987: 340) and Pritchard (1989: 11) Chelonura Temminckii was described by Harlan (1835) on the basis of a specimen collected by Petival in 1835 and several drawings, also by Petival. If indeed Harlan only used the name provided by Troost then MNHN-AC 4540 in the Paris museum may be the holotype (Bour, 1987: 343). However, if Harlan used the name coined by Troost as well as the species description from Troost’s manuscript, then RMNH.RENA 6166 in the Leiden museum should be regarded as the holotype. RMNH.RENA 6166 is a specimen sent by Troost to C.J. Temminck, director of the Leiden mu-seum at that time. King & Burke (1989) also report MNHN-AC A.4540 as the holotype, but mention that Hoogmoed is of the opinion that RMNH.RENA 6166 is the holotype. RMNH records and the original label of the mounted specimen indicate that RMNH.RENA 6166 is the ‘type’ of ‘Macroclemys Temminckii Troost’. The author of the nominal species Chelonura Temminckii therefore is Troost in Harlan, 1835 (fide Hoogmoed, quoted in King & Burke, 1989). The type locality is ‘a tributary stream of the Mississippi, which enters that river above Memphis, in West Tennessee’ [U.S.A.] (Troost in Harlan, 1835: 158). In the RMNH archives there also is a drawing of the head of this species in lateral view in life. Harlan mentions that ‘More than one specimen has been observed’, which might lead to the conclusion that more than one specimen served as the basis for the description, and in that case there would have been several syntypes. It is not clear whether the observations were based on preserved material only, or also included live specimens. However, the fact that this specimen was sent by Troost and the fact that Harlan specifically mentions Troost’s manuscript have convinced us that the RMNH specimen most likely is the holotype of Chelonura Temminckii, and that the specimen in


the Paris museum does not have type status. Duméril & Duméril (1851) described a single specimen, apparently complete, of this species, but its data are not mentioned. Bour (1987) describes a skull in the Paris museum (MNHN-AC A.4540) as the type, but his argumentation is not very convincing and it remains doubtful whether this indeed is a type (see above). According to Fritz & Havaš (2007) the name Chelonura temminckii is a nomen conservandum, see Opinion 660, ICZN 1963. Bour (1987) restricted the type locality to Wolf River, Shelby County, Tennessee, U.S.A. Pictures.― Drawing of side of head in RMNH archives.

Emydidae Lydekker, 1889Emys orbicularis occidentalis Fritz, 1993

*Emys orbicularis occidentalis Fritz, 1993: 136, fig. 8, 10, 12-15.

Paratypes: RMNH.RENA 11371 A-B, 2 juvs. (alc.). Loc.: ‘Tunesia’. Don.: E.C. Stol; RMNH.RENA 15003, 1 ♀ (alc.). Loc.: ‘Ifrane (1650 m), Middle Atlas, Morocco’. Leg.: M.S. Hoogmoed, 23-vi-1967.

Pictures.― Fritz, 1993: 143, pl. 10c (RMNH.RENA 15003).

Geoemydidae Theobald, 1868Cyclemys enigmatica Fritz, Guicking, Auer, Sommer, Wink & Hundsdörfer, 2008

Emys dentata Gray, 1831: errata (part).Emys dhor; Lidth de Jeude, 1898: 6 (part, only Cat. ost. a).Cyclemys oldhamii; Fritz et al., 1997: 198 (part).*Cyclemys enigmatica Fritz et al., 2008: 381.

Paratypes: RMNH.RENA 3838, 1 juv. (alc.) Loc.: ‘Padang, Sumatra’ [Indonesia]. Leg.: [L.] Horner; RMNH.RENA 6066, 6068, two stuffed adult females. Loc.: ‘Sumatra’ [Indonesia], Leg.: [S.] Müller; RMNH.RENA 27828 (Cat. ost. a) skeleton of adult female, Loc.: ‘Java’ [Indonesia]. Leg.: [H.] Kuhl & [J.C.] van Hasselt.

Remarks.― Fritz et al. (2008) incorrectly mention ‘Hoiner’ as collector of RMNH.RENA 3838, this should be Horner (= L. Horner, a member of the Dutch Natuurkundige Commissie). They also incorrectly refer to RMNH.RENA 6088 as a paratype, but this should be corrected to RMNH.RENA 6068. In table 4 on p. 382 the correct number RMNH.RENA 6068 is used. RMNH.RENA 6088 is a Trachemys scripta (Schoepff, 1792) collected by G. Troost in Tennessee. Fritz et al. (1997) identified the above four speci-mens, together with more RMNH material as C. oldhamii Fritz et al. (2008) when de-scribing C. enigmaticus from Sumatra, Borneo, Java and the Malay Peninsula, only con-sidered the above four specimens as paratypes. Fritz et al (1997) considered these four specimens as C. oldhamii Gray, 1863, together with six other RMNH specimens (RMNH.RENA 5003, 6062-65 and 27829) also from Sumatra, Borneo and Java. There is no refer-ence to these last six RMNH specimens mentioned in Fritz et al. (1997) in the Fritz et al. (2008) publication, and thus they cannot be considered type material, although presum-ably they are C. enigmatica as well, as Fritz et al., 1997) did not consider them as C. dentata, the only other species of Cyclemys occurring sympatrically with C. enigmaticus in Sumatra, Borneo, Java and the Malay Peninsula (Fritz et al., 2008).


Cyclemys giebelii Hubrecht, 1881(fig. 1)

*Cyclemys giebelii Hubrecht, 1881: 45.

Syntype: RMNH.RENA 3348, 1 ex. (alc.). Loc.: ‘Borneo’ [Indonesia]. Leg.: C.A.L.M. Schwaner, 1844.

Current name: Notochelys platynota (Gray, 1834). Remarks.― This is the specimen which is fully described by Hubrecht (1881). In his description he also refers to four juvenile specimens from the island Banka, de-scribed by Giebel (1866: 15). These also form part of the type series, but their present whereabouts could not be ascertained. They certainly do not form part of the RMNH collection. Pictures.― Giebel (1866: pl. 3) provides drawings of the syntypes that are not part of the RMNH collection and of which the present whereabouts are not known.

Cyclemys pulchristriata Fritz, Gaulke & Lehr, 1997

*Cyclemys pulchristriata Fritz, Gaulke & Lehr, 1997: 203.

Paratype: RMNH.RENA 4751, 1 juv. (alc.). Loc.: ‘Annam’ [Vietnam]. Leg.: H. Fruhstorfer.

Fig. 1. Notochelys platynota, RMNH.RENA 3348, alcohol preserved syntype of Cyclemys giebelii in original storage bottle. Note the red dot indicating a type specimen and the recent label wrongly suggesting this would be the holotype of N. platynota. Also note the old label with the original name, attached to the outside of the original bottle in the late 19th century, damaged by different climates it was subjected to in the course of time in the storage rooms, but still legible.


Remarks.― Fritz et al. (1997) based their description of C. pulchristiata on material which mostly came from the pet trade. Only material collected by H. Fruhstorfer in Annam was provided with a reliable locality. Material collected by Fruhstorfer is present in museums in Vienna, Leiden, Frankfurt, Stuttgart and Hamburg. In most museums its locality is indicated as ‘Annam’, but the material in Vienna is provided with a more detailed locality: Phuc Son (Annam, Vietnam), and possibly the material in other museums which was provided by Fruhstorfer comes from this locality as well.

Emys borneoensis Schlegel & Müller, 1845(figs 2, 3)

*Emys borneoensis Schlegel & Müller, 1845: 30.Clemmys borneoensis; Strauch, 1865: 87.Batagur borneensis; Hubrecht, 1881: 47.Callagur borneoensis; King & Burke, 1989: 31; Fritz & Havaš, 2007: 213.Batagur borneoensis; Fritz & Havaš, 2007: 367 (Appendix).

Fig. 2. Batagur borneoensis, RMNH.RENA 3296, alcohol preserved juvenile, syntype of Emys borneoensis.

Fig. 3. Batagur borneoensis, RMNH.RENA 6210, stuffed syntype of Emys borneoensis, with original label with information added later in pencil.


Syntypes: RMNH.RENA 3296, 1 ex. (alc.). Loc.: ‘Borneo’ [Indonesia]. Leg.: S. Müller; RMNH.RENA 6210, 1 mounted ex. Loc.: ‘Borneo’ [Indonesia]. Leg.: S. Müller, 1837.

Current name: Batagur borneoensis (Schlegel & Müller, 1845). Remarks.― Schlegel & Müller (1845: 30) mentioned to have seen three specimens of this species deposited in the National Museum of Natural History in Leiden, but only two of them could be located. The third specimen probably was exchanged with one of the large European museums. King & Burke (1989) mention RMNH.RENA 6210 as ‘holotype’. Apparently they were not aware of the existence of RMNH.RENA 3296, and neither of the fact that Schlegel & Müller (1845) based their description on three speci-mens (clearly stating so), without indicating a holotype among them and those three specimens thus are syntypes. King & Burke’s (1989) action cannot be considered a fixa-tion of lectotype as mentioned in article 74.6 of the International Code of Zoological Nomenclature (ICZN, 1999), because it was clear from the beginning that three speci-mens were the basis of the description. Temminck’s ‘Verhandelingen over de Natuurlijke Geschiedenis der Nederlandsche Overzeesche bezittingen, 1839-1844. Zoologie’ was published in parts over a period of 6 years. Husson & Holthuis (1955) provide details on the publication dates of the parts containing reptiles. We here are concerned only with pages 1-36, of which pages 1-28 deal with crocodilians (see below), and pages 29-36 with turtles and tortoises. Plate 4, showing Testudo emys was published on April 24, 1840, pages 29-36, dealing with turtles and tortoises, were published on June 26, 1845. Thus, turtle and tortoise names coined by Schlegel & Müller in Temminck ‘1839-1844’ (Emys borneoensis, Emys subtrijuga, Testudo emys and Testudo forstenii) should be considered having as year of publication 1840 (Testudo emys), or 1845 (Emys borneoensis, Emys subtrijuga and Testudo forstenii). This has no nomenclatural consequences. The accepted name for this species for a long time was Callagur borneoensis (e.g. Fritz & Havaš, 2007), but recent molecular research (Praschag et al., 2007) showed Callagur to be a junior synonym of Batagur. Thus, Fritz & Havaš (2007: 367) had to make a change of the genus name in an Appendix to their checklist. Pictures.― none known.

Emys dentata Gray, 1831

Emys Hasseltii Boie (nomen nudum) in Fitzinger, 1826:45.Emys Dhor Gray, 1831:20.Emys Hasselti; Gray, 1831:20 (part) (nomen nudum).*Emys dentata Gray, 1831: page errata.Emys dhor; Lidth de Jeude: 1898: 6 (part, only Cat. ost. b, c, d).Emys hasseltii; Lidth de Jeude: 1898: 6. Cyclemys dentate; Fritz et al., 1997: 188 (part).Emys hasselti; Fritz et al., 1997: 192 (part).Cyclemys oldhamii; Fritz et al., 1997: 198 (part).

Paralectotypes: RMNH.RENA 6062, 1 mounted ex., Loc.: ‘Java’ [Indonesia], Leg.: [H.] Kuhl & [J.C.] van Hasselt; RMNH.RENA 6063, 1 mounted ex., Loc.: ‘Java’ [Indonesia], Leg.: [H.] Kuhl & [J.C.] van Hasselt; RMNH.RENA 6067, 1 mounted ex., Loc.: ‘Malacca’ [Malaysia] Leg. P. Diard, 1829; RMNH.RENA 40474 (Cat. ost. d), skull, no locality or collector.


Current name: RMNH.RENA 6063, 6067, 40474 Cyclemys dentata (Gray, 1831). RMNH.RENA 6062 Cyclemys enigmatica Fritz, Guicking, Auer, Sommer, Wink & Hundsdörfer, 2008. Remarks.― The name Emys Hasseltii Boie was mentioned by Fitzinger (1826) as a nomen nudum (Fritz & Havaš, 2007) and as Emys Hasselti by Gray (1831) in the synon-ymy of Emys dhor (= dentata) and indicated to be a Boie manuscript name in the Leiden Museum. The name E. Hasseltii was coined by H. Boie (first curator of Herpetology of the RMNH) in his ‘Erpétologie de Java’, which unfortunately was never published. H. Boie prepared a description (with coloured plates) of the material indicated as Emys Hasseltii for publication in his ‘Erpétologie de Java’, which was intended to be pub-lished posthumously, but due to the Belgian insurrection in 1830, the work at the Brus-sels printers was stopped and never started again (Holthuis, 1995; Hoogmoed, 1982). The manuscript and the coloured plates are still in the archives of the Nationaal Natu-urhistorisch Museum in Leiden as a tragic reminder of the collections made by the first explorers of the RMNH that all died too early to be able to see the fruits of their pioneer work in print. Lidth de Jeude (1898), under Emys dhor lists four specimens (three skeletons and a skull), and in the data for Cat. Ost. b and d, he specifically mentions the name Emys hasseltii Boie. Fritz et al. (1997: 192/193) state: ‘Die eindeutige Beschränkung auf nur drei Stücke ist insofern bemerkenswert, als von Gray (1831) angegeben wird, das mit seiner Emys dentata die von ihm im Leidener Museum untersuchte ‘Emys hasselti Boie’ (nomen nudum) identisch sei. Wie wir uns selbst überzeugen konnten, sind im Nationaal Na-tuurhistorisch Museum Leiden eine ganze Reihe von Cyclemys mit diesem Namen etikettiert. Davon lag Gray aber ganz offensichtlich bei der Beschreibung von Emys dentata nur ein einziges Exemplar aus Java vor, welches in den Bestand des Londoner Mu-seums überging und heute die Inventarnummer BMNH 1946.1.22.62 trägt.’ [= The clear restriction to only three specimens is rather remarkable, because Gray (1831) indicates that the ‘Emys hasselti Boie’ (nomen nudum) which he studied in the Leiden museum were identical with his Emys dentata. Like we ourselves could see, in the Nationaal Natuurhistorisch Museum in Leiden there is quite a number of specimens of Cyclemys that are labeled with this name. Of these Gray only had before him for the description of Emys dentata a single specimen from Java, which was transferred to the collection of the London museum and now has the number BMNH 1946.1.22.62]. BMNH 1946.1.22.62 (ex 1828.5.12.1, chosen as lectotype of Emys dhor = E. dentata Gray 1831 by Fritz et al. 1997), was obtained from the Leiden Museum in 1828 and formed part of the series indicated with the name E. Hasselti Boie (Fritz et al., 1997). As Gray (1831: vi-vii) visited the Leiden museum, we may assume that he saw the mate-rial indicated as E. Hasseltii collected by members of the Natuurkundige Commissie, and used those specimens to form his idea of Emys dentata. We therefore consider all specimens in the Leiden museum belonging to either Cyclemys dentata (RMNH.RENA 6063, 6067, 40474) or C. enigmatica (RMNH.RENA 6062), present at the time of Gray’s visit to Leiden and having the name Emys Hasseltii on their original labels, as part of Gray’s type series of E. dentata (which of course also includes the BMNH 1946.1.22.62 lectotype). This is in accordance with the rules of nomenclature (ICZN, 1999) for that period (72.1.1, 72.4.1 and 72.4.1.1), and consequently all four specimens can be consid-ered as paralectotypes. We thus disagree with Fritz et al.’s (1997) opinion that Gray’s


(1831) description of Emys dentata only would be based on three specimens. The RMNH has more material of C. dentata that was collected by members of the Natuurkundige Commissie and was present at the time of Gray’s visit to the Leiden museum, but as these lack any indication of the name E. Hasseltii on the original labels, we have not considered them as paralectotypes of E. dentata Gray 1831. Pictures.― colour plates of Emys Hasseltii in archive RMNH (Erpétologie de Java manuscript of Boie), which at the moment (hopefully temporarily) cannot be found.

Emys subtrijuga Schlegel & Müller, 1845(fig. 4)

Emys trijuga; Temminck & Schlegel, 1834: 64.*Emys subtrijuga Schlegel & Müller, 1845: 30; Hubrecht, 1881: 48.Malayemys subtrijuga; Brophy, 2004: 73.

Lectotype: RMNH.RENA 6082, mounted ex. Loc.: ‘waarschijnlijk in het landschap Bantam’ [= probably in the Bantam region] ‘Java’ [Indonesia]. Leg.: [H.] Kuhl and [J.C.] van Hasselt. Paralectotypes: RMNH.RENA 6084, 6085, two mounted ex. Loc.: ‘waarschijnlijk in het landschap Ban-tam’ [= probably in the Bantam region.] ‘Java’ [Indonesia]. Leg.: [H.] Kuhl and [J.C.] van Hasselt.

Current name: Malayemys subtrijuga (Schlegel & Müller, 1845). Remarks.― Hubrecht (1881) discussed the confusion that reigned about this spe-cies in the early years after its description and categorically stated that the three type specimens were in Leiden and that a specimen in the British Museum (Natural His-tory) received from the Utrecht museum in Holland could not have been one of the types. Brophy (2004) studied the three syntypes and indicated RMNH.RENA 6082 as lectotype. He further discussed the confused nomenclatural history and agreed with Hubrecht (1881) that BMNH 1947.3.4.53 could not be the holotype of Malayemys subtrijuga as had formerly been assumed. Pictures.― colour plates in archive RMNH (Erpétologie de Java manuscript of Boie), which at the moment (hopefully temporarily) cannot be found.

Fig. 4. Malayemys subtrijuga, RMNH.RENA 6682, stuffed lectotype Emys subtrijuga. Note the indication K&vH in the lower left corner of the label, indicating the collectors H. Kuhl and J.C. van Hasselt.


Emys vulgaris Gray, 1831(fig. 5)

Emys vulgaris Gray, 1831: 24 (part).Emys Sigritzii Gray, 1831: 24.Cl. Sigritzii Temminck & Schlegel, 1834: 53.Emys Siegritzii; Fritz & Wischuf, 1997: 248, 249.Mauremys rivulata; Fritz & Wischuf, 1997: 256

Paralectotype: RMNH.RENA 3327 (alc.). Loc. ‘Dalmatia’ [former Yugoslavia]. Leg.: Michahelles.

Current name: RMNH.RENA 3327 Mauremys rivulata (Valenciennes, 1833). Remarks.― The nomenclatural history of RMNH.RENA 3327 is rather confused and complicated. In his description of Emys vulgaris, Gray (1831) mentioned that he had seen a specimen in Leiden [‘Emys Sigritzii: Michaux, MSS (v.Mus. Leyd. Mus.Brit)’], which he considered as belonging to his E. vulgaris (note that Gray on the errata page rectifies Michaux to Michahelles), thus making it a syntype of that name. Gray’s E. vulgaris generally has been considered a synonym of Mauremys leprosa (Schweigger, 1812), but Fritz & Wischuf (1997) pointed out that it was based on a mixture of Mauremys leprosa, M. rivulata and a species of Rhinoclemmys, either R. areolata (Duméril & Bibron, 1851) or R. pulcherrima (Gray, 1856). The name E. vulgaris could constitute a threat to the stability of the last three species names mentioned. In order to avoid nomenclatural upheaval, Fritz & Wischuf (1997) selected the drawing of a juvenile M. leprosa published by Gray (1831 pl. 4, upper drawing) as lectotype of E. vulgaris, thus removing possible problems for the other species names mentioned. They also considered the possibility of RMNH.RENA 3327 being a syntype of Clemmys Sigriz, but discarded this possibility on the basis of Michahelles’ (1829) clear statement about the provenance (Spain) of the specimens of C. Sigriz, which he said were different from specimens from Dalmatia. RMNH.RENA 3327 on the old label on the outside of the bottle in which it is pre-served (still readable in the 1990s, but now illegible) was indicated as being the possible type of Clemmys Siegritzii (sic!, misspelling on label) (Fritz & Wischuf, 1997: 248, 249;

Fig. 5. Mauremys rivulata, RMNH.RENA 3327, alcohol preserved syntype Emys vulgaris. Note 19th cen-tury glass-bulb attached to keep specimen floating upright in bottle.


MSH pers. obs.). This is strengthened by a remark of Temminck & Schlegel (1834: 53) about this name: ‘Mr. Michahelles l’avait déjà décrit d’après les individues, rapports de Dalmatie, et qu’il a bien voulu céder à notre Musée.’ [= Mr. Michahelles already de-scribed it after specimens reported from Dalmatia, and has been so kind to present it to our museum]. In footnote 3 referring to Michahelles’ (1829) publication they use the name Cl. Sigritzii, a name never used by Michahelles, who used C. Sigriz in honour of the collector. However, this taxon has as its type locality ‘Hispaniae meridionalis, venditur in urbibus Hispaniae’ (p. 1300) and not Dalmatia, as suggested by Temminck & Schlegel (1834). Although on the old label a more recent identification as Clemmys leprosa is writ-ten in pencil, the specimen in RMNH.RENA 3327 actually is a halfgrown Mauremys rivulata, a species that occurs near Ragusa (= Dubrovnik) in former Yugoslavia (Fritz & Wischuf, 1997), which is a locality mentioned by Michahelles (1829), being a place where Clemmys caspica [= Mauremys rivulata] had been collected. In the original register-book RMNH.RENA 3327 is registered as: ‘Emys caspica, Dalmatie, Michah.’. Thus there does not seem to be any doubt about the locality and the collector. In Spain the species of Mauremys occurring there is M. leprosa, and most authors have correctly considered Clemmys Sigriz Michahelles, 1829 a synonym of M. leprosa. The indication ‘type’ on the old label of RMNH.RENA 3327 probably was due to the misunderstanding of Tem-minck & Schlegel about the correct identification and allocation of RMNH.RENA 3327.

Emys vulgaris japonica Temminck & Schlegel, 1834(figs 6, 7)

[Emys vulgaris] variété japonaise Temminck & Schlegel, 1834: 54.Emys vulgaris, variété du Japon Temminck & Schlegel, 1834: 77.

Fig. 6. Mauremys japonica, RMNH.RENA 3331, alcohol preserved syntype of Emys vulgaris japonicus.


Fig. 7. Mauremys japonica, RMNH.RENA 3333, alcohol preserved syntypes of Emys vulgaris japonicus in their original storage bottle.


* Emys vulgaris japonica Temminck & Schlegel, 1834: 139.Emys palustris var. Japon Temminck & Schlegel, 1834: pl. 8 figs 1-4; pl. 9.Emys vulgaris japonica; Schlegel, 1844: 126, pl. 41.Emys Japonica; Gray, 1844:19; Gray, 1873: 34.Mauremys japonica; King & Burke, 1989: 42.

Syntypes: RMNH.RENA 3331, 1 ex. (alc.). Loc.: ‘Japan’. Leg.: Ph.F. von Siebold; RMNH.RENA 3332, 1 ex. (alc.). Loc.: ‘Japan’. Leg.: Ph.F. von Siebold; RMNH.RENA 3333, 2 hatchlings (alc.). Loc.: ‘Japan’ Leg.: Ph.F. von Siebold; RMNH.RENA 3334, 1 juv. (alc.). Loc.: ‘Japan’. Leg.: Ph.F. von Siebold; RMNH.RENA 6142, 1 mounted ex. Loc.: ‘Japan’. Leg. Ph.F. von Siebold.

Current name: Mauremys japonica (Temminck & Schlegel, 1834). Remarks.― There is some confusion about the exact year of publication of the part Chelonia of the Fauna Japonica. Wermuth & Mertens (1961) mention 1833, Fritz & Havaš (2007) mention 1835. According to Stejneger (1907: 542-543), the volume Reptilia from the Fauna Japonica was issued in different parts. Chelonii (pp. 1-80, pls. 1-9) ac-cording to him appeared in 1835. Sherborn & Jentink (1895: 149) and Holthuis & Sakai (1970:75, based on Dutch government archives), however, give 1834 as publication date for this section. We here accept 1834 as publication date for the chelonian part of the Fauna Japonica. The name on the original plates is erroneously ‘Emys palustris Var. Japon.’. Tem-minck & Schlegel (1834) acknowledge the mistake in footnote 7 (p. 52): ‘Cette espèce portée par méprise sur notre planche sous le nom d’Emys palustris’ [= Due to a mistake, on our plate this species is mentioned under the name Emys palustris]. Schlegel (1844) mentions the type specimens and says that between 1833 and the moment of writing the RMNH had received from Mr. Bürger drawings of an adult and juvenile specimen made after life in Japan [by the Japanese artist Kawahara Keiga (also called Tojosk by Von Siebold and Bürger)]. Thus it is clear that the specimens depicted on pl. 41 (Schlegel, 1844) are not drawings of any type specimens, and indeed they do not agree with any of the RMNH types listed above. RMNH.RENA 3331 has the hand of the right forelimb mutilated into a clubfoot with three low pyramids of muscular tissue covered by skin. One of the pyramids has a stump of nail, the others not. RMNH.RENA 3333 contains two hatchlings, of which the lower one in the bottle (the one without a floating glass bulb) has been depicted on pl. 8 fig. 1 (Temminck & Schlegel, 1834). Gray (1844; 1873) reports an adult and a half-grown specimen in the collection of the British Museum (Natural History) that were received from the Leiden museum. These specimens might have formed part of the type series. King & Burke (1989) mention RMNH.RENA 3330 and MNHNP 1954 as syntypes of Emys vulgaris japonica, but this is incorrect, as these three specimens are syntypes of Emys vulgaris picta = Chinemys reevesii (see below). Pictures.― Temminck & Schlegel (1834: pl.8, fig.1 (RMNH.RENA 3333), fig. 2 and 3 (RMNH.RENA 3334), fig. 4 (RMNH.RENA ??), pl. 9 (RMNH.RENA 3332); Schlegel (1844: pl. 41).


Emys vulgaris picta Schlegel, 1844(fig. 8)

*Emys vulgaris picta Schlegel, 1844: 127; pl. 42.Emys Japonica; Duméril & Duméril, 1851: 8.

Syntypes: RMNH.RENA 3330 A-B, 2 ex. (alc.). Loc.: ‘Japan’. Leg.: H. Bürger.

Current name: Mauremys reevesii (Gray, 1831). Remarks.― The publication date on the first page of Schlegel’s Abbildungen is 1837-1844, and the work was issued in parts. See Stejneger’s Herpetology of Japan (1907: 540) for the exact dates of publication. Schlegel (1844) affirmed that three specimens of this species were deposited in the National Museum of Natural History in Leiden by Bürger, but only two of them have been located in the RMNH collections. Duméril & Duméril (1851) mention that the Paris museum has one specimen of this species, received from the Leiden museum. This might very well be the ‘missing’ third specimen mentioned by Schlegel (1844) and thus can be regarded as one of the syntypes (MNHNP 1954). At the time there was a lively contact between the Paris and Leiden museums, with many specimens being ex-changed, and G. Bibron spending time in the Leiden collection. Despite using the name Emys Japonica for their specimen, Duméril & Duméril (1851) further in the text clearly refer to ‘E. vulgaris japonica picta Schlegel, Abbildung. Neuer Amphib., p. 127, pl. 42’. It should be noted that Schlegel (1844) never used the name Emys vulgaris japonica picta, as mentioned by Duméril & Duméril (1851) for this taxon, just Emys vulgaris picta The drawing on pl. 42 (Schlegel, 1844) sent by Bürger is of an adult specimen in life, [made in Japan by Kawahara Keiga] and does not agree with the adult specimen in RMNH.RENA 3330 (the pattern of lines on the neck and side of the head is clearly dif-ferent). Pictures.― Schlegel (1844): pl. 42.

Fig. 8. Mauremys reevesii, RMNH.RENA 3030 A-B, alcohol preserved syntypes of Emys vulgaris picta. Note the wooden sticks used to keep the specimens in position, apparently commonly used in the early 19th century to pose specimens.


Geoemyda spengleri sinensis Fan, 1931

*Geoemyda spengleri sinensis Fan, 1931: 146.

Paratype: RMNH.RENA 5887, 1ex. (alc.). Loc.: ‘Loshiang’ [Yao-shan, Kwangsi, South China]. Leg.: S.S. Sin. Don.: Sun Yat-Sen University, Guangzhou, P.R. China.

Current name: Geoemyda spengleri (Gmelin, 1789). Pictures.― None known.

† Hardella isoclina Dubois, 1908

*Hardella isoclina Dubois, 1908:1270.Clemmys ? isoclina; Williams, 1957: 235.

Holotype: RMNH Dub. 2722 (carapace and plastron): Pleistocene, Trinil Beds, Kedoeng Panas, Java, Indonesia.

Current name: † Mauremys? isoclina (Dubois, 1908). As Fritz & Havaš (2007) do not include fossil species, the present identification is based on ‘circumstantial evidence’ as provided by Williams (1957) who considered this species most closely related to Clemmys mutica (see below). Remarks.― Dubois (1908) only briefly diagnosed this taxon. Williams (1957) stud-ied the type and referred material from the RMNH Dubois collection and came to the conclusion that ‘The Trinil form appears to resemble most closely the living species mutica from southern China, Formosa, Hainan, and Japan, and since mutica is currently though questionably assigned to the genus Clemmys it will be convenient for the present to refer the Dubois species to Clemmys with a query’. Williams (1957) provided an ex-tensive description and several photographs and drawings of the holotype. Chelonian systematics recently has been in turmoil, especially concerning South-east Asian species. Clemmys nowadays only contains a single, small species and its dis-tribution is restricted to eastern North America. Clemmys mutica now is considered a member of Mauremys (Fritz & Havaš, 2007), and became Mauremys mutica. Mauremys is a genus occurring in southern Europe, northern Africa, the Middle East, Southeast and East Asia (China, Taiwan, Vietnam, Korea, and Japan). The presence of a fossil Mauremys on Java would constitute a considerable extension of its distribution area, but would not be too farfetched. However, the present identification is only based on the nomenclatorial changes involving Mauremys mutica, and new study of the holotype would be very desirable to come to a definite conclusion. Jaekel (1911) describing fossil remains from the Trinil beds does not mention Dubois’s Hardella isoclina, but only men-tions two species of Batagur, which also belong to the family Geoemydae. Pictures.― Williams (1957).

Testudinidae Gray, 1825Testudo dussumieri Gray, 1831

*Test. Dussumieri; Gray, 1831a:3 (nomen nudum); Gray, 1831b: 9; Temminck & Schlegel, 1834: 75.Testudo gigantean; Hubrecht, 1881: 43.


Testudo dussumieri Bour, 1984: 171; Bour, 2006: 20-22; Pritchard, 1986: 532; Frazier, 2009: 39.Dipsochelys dussumieri; Gerlach, 2004: 67; Bour, 2006: 21, 22; Grünewald, 2009b: 136.[Testudo] dussumieri; Cheke, 2009: 175.

Lectotype: RMNH.RENA 3231, 1 juv. (alc.) Loc.: ‘Île Aldabra près de Madagascar’ [Aldabra Islands, Seychelles]. Leg.: J.-J. Dussumier.

Current name: Dipsochelys dussumieri (Gray, 1831) Remarks.― Hoogmoed & Crumly (1984) did not mention this specimen because at the time it had not yet been recognised as a type specimen. Bour (1984) was the first to report it as ‘le type’ after Temminck & Schlegel (1834) and Hubrecht (1881) had men-tioned this specimen without reference to its status. According to Temminck & Schlegel (1834) this specimen from ‘Aldebra’ (sic!) was received under the name T. Dussumieri from the Paris museum, thus not directly from Dussumier, but as a donation or an ex-change from the Paris Museum. This is the only reference about the acquisition of this specimen, as neither in Paris, nor in the Leiden archives any documentation about the way it was obtained could be found (Bour, 2006: 22). Gray (1831b) provided a short description of a juvenile specimen, which he saw in the Leiden museum. He attributed the name to Schlegel (but see statement above by Temminck & Schlegel (1834) from which it is clear that the name was already provided by the Paris museum), but as Schlegel (or anybody else) never published this name, Gray becomes the author. Gray (1831a) described Testudo dussumieri in the synonymy of Testudo indica as a nomen nudum, but Gray (1831b) provided a short description . Bour (2006) designated the specimen as the lectotype, as Gray (1831b) also referred to a fig-ure of another specimen and thus based himself on a series of two specimens. However, the statement by Temminck & Schlegel (1834) mentioned above shows that references in the literature to Testudo dussumieri Schlegel (in Gray) (e.g. Gray, 1831a, 1831b: 9; Fitzinger, 1835: 122; Fritz & Havaš, 2007: 265) are not correct. Schlegel did not coin the name, neither in a manuscript, nor as a label name, apparently somebody in Paris coined the name. Gray (1831b: vii) following the agreement with Temminck, mentioned that he saw a specimen with this name on its label in the Leiden museum, but errone-ously assumed that this label name had been given by Schlegel, which according to Schlegel’s own words (see above) was not true. After Bour (1984) and Pritchard (1986) published their papers doubting the identity of the holotype of Testudo gigantea Schweigger, 1812 (at that time considered lost) there has been a vivid discussion concerning the correct name, both generic and specific for the Aldabra tortoise. Frazier (2006), just for nomenclatural, not for taxonomic reasons, designated a neotype, with the intention to stabilise the name. However, Bour (2006) reported the rediscovery of Schweigger’s holotype, which clearly is a specimen of Geochelone denticulata (Linnaeus, 1766) (as already hypothesized by Pritchard (1986)). This re-discovery of the holotype clearly shows that T. gigantea Scheigger, 1812 is a junior synonym of T. denticulata and makes the designation of the neotype void. Bour (1984: 171, footnote 1) mentioned the existence of RMNH.RENA 3231 and provisionally con-sidered it a ‘nomen oblitum’. Bour (2006) designated this specimen as lectotype of Testudo dussumieri and considered it available. Frazier (2009) appealed to the International Commisision of Zoological Nomenclature (Case 3463) to maintain the specific name


gigantea, recognise the neotype he designated and suppress T. dussumieri. Bour & Pritchard (2009) made a very clear defence for disregarding the neotype of T. gigantea and accepting the name Dipsochely dussumieri, which has been in regular use since the 1990’s for the Aladabra tortoise. Although the ICZN has not yet given its opinion on which scientific name should be used for the Aldabra tortoise, we here follow Bour & Pritchard (2009) in recognising that, due to the new situation with the holotype surfac-ing, the neotype designation is no longer valid and that the name T. dussumieri becomes available, based on the perfect juvenile specimen RMNH.RENA 3231, completely pre-served in alcohol, that without any doubt belongs to the Aldabra tortoise and has a good locality. The specimen was reported to come from ‘Aldebra’ (sic!) by Temminck & Schlegel (1834), and this same locality (Île Aldabra prés de Madagascar) [= Aldabra is-land, near Madagascar] also appears in the old handwritten register of the RMNH and on the original handwritten label of RMNH.RENA 3231. Matyot (2009) doubts whether Dussumier ever visited Aldabra, but Bour et al. (2010) and Cheke (2010) showed that Dussumier, even if he would not have visited Aldabra, easily could have obtained ma-terial from there. However, this all remains speculative, the data on the label of RMNH.RENA 3231 are the only firm data that exist and have to be accepted as correct. It should be noted (Dubois et al, 2010) that despite all the vehement discussions on this issue, only three professional herpetologists (R. Bour, M.S. Hoogmoed, P.C.H. Pritchard) have ever studied RMNH.RENA 3231 and all three came to the conclusion that RMNH.RENA 3231 is a juvenile Aldabra tortoise. Pictures.― Gerlach, 2004: 68 (dorsal, lateral and ventral view); Bour, 2006: 21, fig. 3 (lateral, dorsal and ventral view); Grünewald 2009b: 137 (dorsal and ventral view), 138 Lateral view and detail head), 139-141 (labels).

Testudo emys Schlegel & Müller, 1840(fig. 9)

*Testudo emys Schlegel & Müller, 1840: pl. 4; Schlegel & Müller, 1845: 34; Lidth de Jeude, 1896: 197, pls. 5, 6; Lidth de Jeude, 1898: 4.

Fig. 9. Manouria emys, RMNH.RENA 17967, skull of syntype of Testudo emys.


Testudo emydoides Duméril & Duméril, 1851: 4.Manouria emys; Hoogmoed & Crumly, 1984: 251; King & Burke, 1989: 92.

Lectotype: RMNH.RENA 3808, 1 ex. (alc.). Loc.: ‘Sumatra’ [Indonesia]. Leg.: S. Müller. Paralectotypes: RMNH.RENA 6005, 1 mounted ex., RMNH.RENA 6030, 1 mounted ex., both: ‘Batang Singalang, Sumatra’ [Indonesia]. Leg.: S. Müller. RMNH.RENA 17967 (Cat. ost. a, 41 cm), 1 skeleton. Loc.: ‘Batang-Singalang’ [Indonesia]. Leg.: S. Müller.

Current name: Manouria emys emys (Schlegel & Müller, 1840). Remarks.― Due to the complicated publication history of Temminck’s ‘Verhande-lingen over de Natuurlijke Geschiedenis der Nederlandsche Overzeesche bezittingen, 1839-1844. Zoologie’ (see under Emys borneoensis), with plates and parts appearing in different fascicules in different years, the publication date of Testudo emys must be con-sidered to be (April 24) 1840, the year in which plate 4, on which the species is pictured with its name, appeared (Husson & Holthuis, 1955). Only on June 26, 1845 the complete description of Testudo emys appeared (Husson & Holthuis, 1955). Schlegel & Müller (1840, 1845) based their description on a series of six specimens, which consequently were the syntypes. Four of these are still present in Leiden. One was exchanged with the Museum d’Histoire Naturelle, Paris (MNHN 9422) (Duméril & Duméril, 1851; Hoogmoed & Crumly, 1984). This Paris specimen is another paralectotype of Testudo emys, received from the Leiden Museum, collected by S. Müller at the river Aneh, Su-matra and it also is the holotype of Testudo emydoides Duméril & Duméril, 1851. Schlegel & Müller (1845: 36) give as locality for ‘three or four’ of the specimens: ‘Sumatra (…) aan de zuiderzijde van den Goenong Singalang (…) rivier Aneh’ [Su-matra, south side of Gunung Singalang, (…) Aneh river]. They also received some other (‘eenigen’) specimens from ‘de voorbergen beoosten Padang’ [low mountains east of Padang, Sumatra]. Hoogmoed & Crumly (1984: 251) designated RMNH.RENA 3808 as lectotype. The lectotype RMNH.RENA 3808 is the specimen depicted by Müller & Schlegel (1840, pl. 4). Lidth de Jeude (1896) discussed the Leiden specimens of Manouria emys (indicating specimens with letters that have no relation to the letters used in his ‘Catalogue Os-téologique’), among which four syntypes, extensively and provided pictures of plastra and a skull of RMNH.RENA 40186 (cat. ost b (not a syntype)). It seems useful to provide additional data on the material mentioned by him. His specimen ‘a’ is RMNH.RENA 3808, ‘b’ is RMNH.RENA 6005, ‘c’ is RMNH.RENA 6030, ‘d’ is RMNH.RENA 17967 (cat. ost. a), ‘e’ was a recently acquired specimen (January 1896, Padang, Zool. Garden Rot-terdam) RMNH.RENA 40186 (cat. ost. b), and ‘f’ was a specimen at the time still alive in the Zoological Garden Rotterdam. Thus, Lidth de Jeude’s (1896) ‘a’, ‘b’, ‘c’ and ‘d’ were the syntypes of the species. RMNH.RENA 17967 is a skeleton that still has the horny scales attached to the carapace. The skull and lower jaw, the cervical vertebrae and the bones of the right forelimb have become detached from the skeleton and are stored in a separate cardboard box. Pictures.― Schlegel & Müller, 1840: pl. 4, figs 1-5 (RMNH.RENA 3808); Lidth de Jeude, 1896: pls. 5 fig. 1 (RMNH.RENA 17967), fig. 3 (RMNH.RENA 6030); Hoogmoed & Crumly, 1984: 252 (fig. 5), 253 (fig. 6 = RMNH.RENA 3808).


Testudo Forstenii Schlegel & Müller, 1845

*Testudo Forstenii Schlegel & Müller, 1845: 30.Indotestudo forstenii; Vervoort, 1981: 246; Hoogmoed & Crumly, 1984: 243; King & Burke, 1989: 89.

Holotype: RMNH.RENA 3811, 1 ex. (alc.)(carapace 25 cm). Loc.: ‘Gilolo’ [Halmahera, Indonesia]. Leg.: E.A. Forsten.

Current name: Indotestudo forstenii (Schlegel & Müller, 1845). Remarks.― As can be seen in the pictures provided by Vervoort (1981) and by Hoogmoed & Crumly (1984) (actually the dorsal and ventral views are the same pho-tos) a number of marginal scutes had already become detached from the bony carapace at that time. At the moment of writing more scutes (including costals) have become detached and parts of them are still in the container with the specimen. For further com-ments on this specimen see Hoogmoed & Crumly (1984) who also provide measure-ments. Pictures.― Vervoort, 1981: 246 (fig. 9, dorsal and ventral view); Hoogmoed & Crumly, 1984: 246 (fig. 2, dorsal, ventral and lateral view).

Testudo Strauchi Lidth de Jeude, 1893

*Testudo Strauchi Lidth de Jeude, 1893: 312, pl. 9.Psammobates geometricus; Hoogmoed & Crumly, 1984: 257.

Holotype: RMNH.RENA 6011, 1 mounted ex. Loc.: ‘Cape of Good Hope’ [South Africa]. Leg.: [H.] Kuhl and [J.C.] van Hasselt.

Current name: Psammobates geometricus (Linnaeus, 1758). Remarks.― The specimen was complete in 1893 (see Lidth de Jeude, 1893: pl. 9). The left hind leg of this specimen is now missing, as already shown in Hoogmoed & Crumly (1984: fig. 8), and the right front leg now is detached from the body, but still kept in the cardboard box in which the specimen is kept. The specimen also shows some paint marks on the plastron, probably caused by former mounting on a freshly painted wooden pedestal. Pictures.― Lidth de Jeude, 1893, pl. 9; Hoogmoed & Crumly, 1984: 256 (fig. 8).

Testudo indica Vosmaeri Suckow, 1798

Testudo indica. Vosmaeri - Schoepff, 1792: 103 and pl. 22 middle and lower figures; Schoepff, 1801: 120 and pl. 22 (middle and lower figure).

*Testudo indica Vosmaeri Suckow, 1798: 57. Testudo Vosmaeri; Fitzinger, 1826: 44; Günther, 1877: 53; Hubrecht, 1881: 41; Vaillant, 1893: 18. Testudo indica; Temminck & Schlegel, 1834: 74.Testudo vosmaeri; Lidth de Jeude, 1898: 4.Geochelone vosmaeri; Hoogmoed & Crumly, 1984: 242; Hoogmoed, 1999: 57.Cylindraspis vosmaeri; Gerlach, 2004: 103.

Holotype: RMNH.RENA 6001 (Cat. ost. a), 1 ♂, shell. Loc.: ‘Africa, Promontorio bonae spei’ [Cape of Good Hope, South Africa]. Ex.: Collection Prince d’Orange [= collection of Stadtholder].


Current name: Cylindraspis vosmaeri (Suckow, 1798). Remarks.― Temminck & Schlegel (1834) indicated that the species did not occur in the Cape of Good Hope, and Günther (1877) specified that this specimen probably came from the island Rodriguez, which according to Fritz & Havaš (2007) constituted a type locality designation. The history of this specimen was discussed by Temminck & Sch-legel (1834: 74-75), Hubrecht (1881) and Hoogmoed & Crumly (1984), who also pro-vided measurements of the specimen. RMNH.RENA 6001 is a complete carapace and plastron joined, with the horny scutes still attached. The plastron is concave, so this was a male. Based on a remark by Temminck & Schlegel (1834: 75), Hoogmoed & Crumly (1984) noted that this specimen also would be a syntype of Chersine retusa Merrem, 1820, but reconsidering Merrem’s (1820) text, we now are of the opinion that this was not the case and that Merrem’s name seems just a replacement name for Testudo indica Schneider, 1783 (Fritz & Havaš, 2007). Vaillant (1893) also discussed this specimen and correctly noted ‘Le plus intéressant comme étant le type de l’espèce est la carapace que décrivait Schoepff en 1792…’ [= The most interesting for being the type of the species is the carapace which Schoepff described in 1792.]. Gerlach (2004: 103) apparently consid-ers this a lectotype indication and mentions MNHNP 1883-558 as paralectotype. How-ever, this is erroneous, as Schoepff (1792) and Suckow (1798) only based themselves on one specimen, viz. RMNH.RENA 6001, which by definition thus is the holotype. MNHNP 1883-558 at the time of describing Testudo vosmaeri was not taken into account and just is another specimen of this taxon, but not a type. Note that the original description and figure by Schoepff (1792) mention the name ‘.Vosmaeri’ preceded by a dot, which in this case means Testudo indica sensu Vosmaer. Thus, Schoepff (1792) did not propose the specific or subspecific name ‘vosmaeri’ for this taxon, he was just relaying Vosmaer´s information on the specimen (Hoogmoed & Crumly 1984). The first one to officially coin the name Testudo indica Vosmaeri was Suck-ow (1798), who thus becomes the author of the taxon. The species is considered extinct since about 1795 (Fritz & Havaš, 2007), apparently shortly after the RMNH specimen reached the Netherlands. However, Pritchard (1986) citing Bour mentions around 1804 as the date of disappearance of this species. As indicated by the text on the label, this specimen originally formed part of the Cabinet of the Stadtholder, which was donated to the collection of Leiden University, which in 1820 formed one of the constituent parts of the collections of the Rijksmuseum van Natuurlijke Historie. It is not clear whether this specimen was stolen by the French in 1795 and transported to Paris and returned to the Netherlands in 1815 by the efforts of Prof. Dr. S.J. Brugmans (Holthuis, 1995), or whether it formed part of the collection of the Stadtholder that was overlooked by the French and stayed in Holland through-out the French occupation (Boeseman, 1970). Pictures.― Schoepff, 1792: pl. 22 middle and lower figures; Schoepff, 1801: pl. 22 middle and lower figure; Hoogmoed & Crumly, 1984: 246 (fig. 2): Hoek Ostende et al., 1999: 57 (picture Purcell of carapace in right lateral view, and detail of last vertebral scale).


Trionychidae Fitzinger, 1826Trionyx stellatus var. japonicus Temminck & Schlegel, 1834

(fig. 10)

*Trionyx stellatus var. japonicus Temminck & Schlegel, 1834: 32; pl. 5 fig. 7; pl. 7, figs 1, 2; Schlegel, 1844: 108, pl. 31.

Syntypes: RMNH.RENA 3259, RMNH.RENA 3264, 2 ex. (alc.). ‘Japon’. Leg.: Ph.F. von Siebold.

Current name: Pelodiscus sinensis (Wiegmann, 1834). Remarks.― Several years of publication have been associated with the volume Reptilia of the Fauna Japonica. See the remarks section under Emys vulgaris japonica for our decision to accept 1834 as publication date for the chelonian part of the Fauna Japonica. Temminck & Schlegel (1834: 34) stated to have received six specimens from Von Siebold, all of them preserved in alcohol (‘les six sujets envoyés du Japon par Mr. Von Siebold et conservés à l’esprit de vin.’) [= the six specimens sent from Japan by Mr. Von Siebold and preserved in alcohol.]. Only two syntypes have been located in the Na-tional Museum of Natural History in Leiden. Schlegel (1844: 109) states that since de-scribing this taxon the Leiden museum had received a drawing and ‘eine vollständige Reihe in Weingeist aufbewahrter Individuen’ [= a complete series of specimens pre-served in alcohol]. Based on these additional specimens he provides a detailed addition to the original description. The drawing on pl. 31 (Schlegel, 1844) sent by Bürger apparently is of an adult

Fig. 10. Pelodiscus sinensis (species complex), RMNH.RENA 3264, alcohol preserved syntype of Trionyx stellatus var. japonicus. Note the wooden sticks used to keep the specimen in position, apparently com-monly used in the early 19th century to pose specimens.


specimen in life [made in Japan by Kawahara Keiga], and as expected does not agree with any of the specimens in the RMNH collection. We have provisionally identified this taxon as P. sinensis, being aware of the recent publication by Fritz et al. (2010) from which it seems to be clear that this is a species complex that in future may need further subdivision. The Japanese population does not seem to be natural but is derived from introduced specimens (Fritz et al., 2010) of which the provenance apparently is not (yet) known. For the time being it seems best to associ-ate Trionyx stellatus var. japonicus with P. sinensis and hope that in the future a more precise synonymization will be possible. Pictures.― Temminck & Schlegel, 1834: pl. 5 fig. 7; pl. 7, figs 1, 2; Schlegel, 1844: pl. 31.

TESTUDINESPLEURODIRA

Chelidae Gray, 1831Batrachemys heliostemma McCord, Joseph-Ouni & Lamar, 2001

*Batrachemys heliostemma McCord, Joseph-Ouni & Lamar, 2001: 734, figs 1, 3-6, 8.Mesoclemmys heliostemma Grünewald, 2009a: 16.

Paratypes: RMNH.RENA 31998-31999, 2 juvs. (alc.). Loc.: ‘Callao, on the north bank of the Río Tapiche [5°21’S, 47°9’W] near the mouth of the Río Blanco, a tributary of the Río Ucayali, Loreto, Perú’. Leg.: F. Medem.

Current name: Mesoclemmys heliostemma (McCord, Joseph-Ouni & Lamar, 2001) Pictures.― RMNH.RENA 31998 in McCord et al. 2001: 735, fig. 6D and E and in Grünewald 2009a: 17 (dorsal and ventral view), 18 (lateral view and detail head). RMNH.RENA 31999 in Grünewald 2009a: 19 (dorsal and ventral view), 20 (lateral view and detail head).

Chelodina mccordi Rhodin, 1994

Chelodina novae guinea; Kate, 1894: 688. Chelodina novaeguineae; Lidth de Jeude, 1895: 120; Rooij, 1915: 315 (part), 350.Chelodina novae guinea; Lidth de Jeude, 1898: 10.*Chelodina mccordi Rhodin, 1994: 4.

Paratype: RMNH.RENA 10187 (Cat. ost. C. novaeguineae), 1 skeleton, 19 cm. Loc.: ‘Rotti’ [= Roti Island, Indonesia]. Leg.: H.F.C. ten Kate.

Remarks.― Lidth de Jeude (1895: 120) considered three turtles collected by Dr. H. [F.C.] ten Kate on Roti as Chelodina novaeguineae. From his description it is clear that the museum received three specimens, two small ones and a larger one. The larger speci-men mentioned is RMNH.RENA 10187 (a skeleton with the horny plates attached to the carapace and plastron), the two smaller ones, who were only considered as referred material in the description of C. mccordi, are RMNH.RENA 4349.


Dr. L.D. Brongersma, former herpetologist and director of the RMNH. in the 1950’s already had recognized these three specimens as belonging to a new species, which he intended to describe as C. rottiensis, and the specimens were marked as such, even bear-ing labels of holotype and paratypes. However, this name was never published and is superseded by Rhodin’s name. In the description Rhodin (1994) duly mentions the fact that Dr. L.D. Brongersma had recognized the RMNH specimens as a new species and had relinquished claims to the original description. Rhodin (1994) described the species based on the holotype (MCZ 176730) and on three paratypes (MCZ 176731, 176732 and RMNH.RENA 10187). He also mentioned 14 referred specimens (one of the specimens in RMNH.RENA 4349, AGJR 365-7, 369, 448-9, 452-7, 460) [AGJR = Anders G.J. Rhodin private collection eventually to be deposited in MCZ]. It is not clear why the 14 referred specimens are not considered paratypes as well, because the author does not provide any reason why they should be considered as referred material only. On the contrary, all morphometric data of carapace and plas-tron are based on all specimens available. For skull morphometrics only the holotype and the three paratypes were used. It is interesting to note that only the RMNH speci-mens with certainty were obtained on Roti Island by Dr. H.F.C. ten Kate and that his itinerary (Kate, 1894) served to restrict the type locality to ‘Danau Naloek, near Busalang-ga, ca. 11 km northeast of Tudameda and ca. 8 km southwest of Ba´a, elevation ca. 115 m, southwestern Roti Island (10°48´S, 123°00´E). East Nusa Tenggara Province, Indone-sia’ [Rhodin, 1994: 10]). It should be noted that this is a locality where Dr. ten Kate bought (not observed as stated by Rhodin, 1994) a small specimen of ‘Chelodina novae guineae’ and his two small specimens now are preserved (in good condition) as RMNH.RENA 4349, which strangely enough was considered referred material only by Rhodin. All other material in MCZ and AJGR (including the holotype) was ‘purchased from na-tive villagers by Frank Yuwono in Kupang, western Timor, originally collected on Roti Island’ (Rhodin, 1994) and thus at best is of questionable origin. The referred RMNH.RENA 4349 juvenile specimen has a carapace length of 99.5 mm, a midline plastron length of 76 mm and a total plastron length of 81 mm. Pictures.― Rhodin (1994): fig. 4 skull of RMNH.RENA 10187 in dorsal, ventral and lateral view; fig. 5 plastron and carapace RMNH.RENA 10187.

Podocnemididae Cope, 1868Emys Amazonica Spix, 1824

(fig. 11)

*Emys Amazonica Spix, 1824: 1; pl. I; pl. II, figs 1-3; Franzen & Glaw, 2007: 207.Emys expansa; Temminck & Schlegel, 1834: 48.Podocnemis expansa; Hoogmoed & Gruber, 1983: 342.

Paralectotype: RMNH.RENA 3294, 1 juv. (alc.). Loc.: ‘Macañon’ [probably Marañon, see Hoogmoed & Gruber, 1983] [Brazil]. Leg.: J.B. von Spix.

Current name: Podocnemis expansa (Schweigger, 1812). Remarks.― The lectotype of Emys Amazonica Spix was selected by Hoogmoed & Gruber (1983: 343). It is housed in the Zoologische Staatssammlung München (ZSM


2446/0/1), together with a number of paralectotypes (ZMSH 7/0-14/0, 2446/0/2-7, 2447/0/1-4, 2730/0 and 3095/0). The numbers in ZSM differ slightly from those provided by Hoogmoed & Gruber (1983) and now follow Franzen & Glaw (2007), who provided the actual numbering system of ZSM. The type locality given by Spix (1824: 2) is ‘Habitat in fluvio Solimoens et confluentibus Javary, Rio Branco’ [Habitat in Rio Solimões and at the confluence of Javary and Rio Branco], which actually is a composite of two localities which are separated by hundreds of kilometers: Rio Solimões, in Amazonas State and the confluence of the Javari and Rio Branco in northern Brazil, Roraima State. Temminck & Schlegel (1834:48) refer to this species and say ‘Le Musée des Pays-Bas doit à celui de Munich, les individus qu’il possède’ [= The museum of Holland received from that in Munich the specimens it has]. From this text it can be concluded that the Leiden museum originally had more than one specimen of this species, but at the mo-ment only RMNH.RENA 3294 is present and no other specimens that would qualify as Spix material. Pictures.― none known.

Emys macrocephala Spix, 1824

*Emys macrocephala Spix, 1824: 5; pl. IV; Franzen & Glaw, 2007: 207.Emys dumeriliana; Temminck & Schlegel, 1834: 48.Peltocephalus tracaxa; Hoogmoed & Gruber, 1983: 345.

Lectotype: RMNH.RENA 6164, 1 mounted ex. Loc.: ‘Rio Solimoëns’ [Brazil]. Leg.: J.B. von Spix.

Current name: Peltocephalus dumerilianus (Schweigger, 1812). Remarks.― Temminck & Schlegel (1834: 48) briefly discuss the RMNH specimen and synonymise it with Emys dumeriliana Schweigger, 1812. Hoogmoed & Gruber (1983: 347) discussed the nomenclature of this species and selected the lectotype. Two paralec-totypes (skulls) are in the Zoologische Staatssammlung München (ZSM 15/0, 17/0). The type locality given by Spix (1824: 5) is ‘Airon ad ripam fluminis Yau, confluentis Rio

Fig. 11. Podocnemis expansa, RMNH.RENA 3294, alcohol preserved, recently hatched juvenile, syntype of Emys amazonica.


Negro’ [Airon at the banks of the river Yau, at the confluence with the Rio Negro] (= Airon on the confluence of the Rio Jaú and Rio Negro, north of Manaus, Brazil). Ac-cording to Hoogmoed & Gruber (1983: 347), the locality on the RMNH label is likely to be erroneous. Hoogmoed & Gruber (1983) already doubted whether the specimen depicted by Spix (1824) is the RMNH lectotype. They noted differences in the position of head and legs and in the borders of scales. RMNH.RENA 6164 has the head dis-tinctly more extended than the animal depicted on pl. 4 (Spix, 1824) and we think the differences are such that the specimen depicted must have been another individual than RMNH.RENA 6164. Pictures.― Spix, 1824, pl. 4 (another specimen than the RMNH lectotype).

CROCODYLIA

Crocodylidae Gray, 1825Crocodilus biporcatus raninus Müller & Schlegel, 1841

Crocodilus biporcatus; Müller, 1838:75 (part).*Crocodilus biporcatus raninus Müller & Schlegel, 1839: pl. 3, figs 7, 8; Müller & Schlegel, 1841: 26.Crocodilus porosus – Gray, 1844: 58 (part); Strauch, 1866 (part); Lidth de Jeude, 1898: 14 (part).Crocodilus palustris; Strauch, 1866 (part).Crocodylus raninus; Ross, 1990: 960; Ross, 1992: 401.

Lectotype: RMNH.RENA 3219, 1 juv. (alc.). Loc.: ‘Pontianak, Borneo’. Leg.: P. Diard.Paralectotypes: RMNH.RENA 7939 + RMNH.RENA 21695 1 juv., respectively dry skin and skull + lower jaw (of the same specimen). Loc.: ‘op Java (…), omstreken van Tjikao’ [near Kao Tji, Jawa Barat, Indone-sia]. Leg.: [H.] Boie and [H. C.] Macklot; RMNH.RENA 37489 (formerly Cat. ost. n, 68 cm), 1 skull. Loc.: ‘op Borneo (…) tot ver in de binnenland toe’ [interior of Borneo]. Leg.: S. Müller, 1836; RMNH.RENA 37493 (formerly Cat. ost. p, 46 cm), 1 skull. Loc.: ‘Tribuary of the Banjer River, Borneo’. Leg.: S. Müller, 1836.

Current name (according to Ross, 1990, 1992): RMNH.RENA 3219 (lectotype) and RMNH.RENA 37493 (paralectotype) Crocodylus raninus Müller & Schlegel, 1841. RMNH.RENA 37489 (paralectotype) Crocodylus porosus Schneider, 1801. RMNH.RENA 7939 + 21695 (paralectotype) and 1 juv. SMF 8090 (paralectotype) Crocodylus siamensis Schneider, 1801. Remarks.― Ross (1992) recognized three different species in the type series. RMNH.RENA 3219 and RMNH.RENA 37493 (Cat. ost. p) are the only specimens that belong to Crocodylus raninus. RMNH.RENA 37489 (Cat. ost. n) is a skull of C. porosus Schneider, 1801 and RMNH.RENA 21695 (+ RMNH.RENA 7939), as well as a syntype now in the Forschungsinstitut und Natur-Museum Senckenberg in Frankfurt (SMF 8090, juvenile, ex RMNH) belong to C. siamensis Schneider, 1801. Ross (1990: 960) revived Crocodylus raninus Müller & Schlegel, 1844 as a distinct species. In a following publication, he se-lected RMNH.RENA 3219 as lectotype, and RMNH.RENA 37493, RMNH.RENA 37489 (Cat. Ost. n), RMNH.RENA 21695, RMNH.RENA 7939 and SMF 8090 as paralectotypes (Ross, 1992: 401). After the publication of both articles by Ross, the specimens referred to as RMNH


‘n’ and RMNH ‘p’ have been renumbered (now RMNH.RENA 37489 and RMNH.RENA 37493). Ross (1992) provided new measurement for the skulls RMNH.RENA 37489 (64.5 cm) and RMNH.RENA 37493 (45 cm), which differ slightly from the sizes mentioned by Lidth de Jeude (1898). Lidth de Jeude (1898) under Crocodilus porosus, indicated both skulls Cat. Ost. n (= RMNH.RENA 37489) and Cat. Ost. p (= RMNH.RENA 37493) as: ‘ race à museau obtus’ [= race with the blunt snout] and gave lengths of respectively 68 cm and 46 cm. RMNH.RENA 7939 (skin) and 21695 (skull + lower jaw) belong to the same juvenile specimen of Crocodylus siamensis, that apparently originally was mounted on a wooden pedestal (still present) until at least 1898, because the skull RMNH.RENA 21695 is not mentioned by Lidth de Jeude (1898) in the Catalogue Ostéologique. Apparently at a later date the skull and the stuffing were removed from the skin and skin and skull ended up with different registration numbers. It is not clear what happened to the rest of the skeleton, but no matching skeleton could be found in the RMNH collections. The dried skin of the specimen referred to as RMNH.RENA 21695 by Ross (1990, 1992) actu-ally is numbered as RMNH.RENA 7939, while the skull of the same specimen remains RMNH.RENA 21695. Temminck’s ‘Verhandelingen over de natuurlijke geschiedenis der Nederlandsche Overzeesche bezittingen, 1839-1844. Zoologie’, as mentioned above, was published in parts over a period of 6 years. Husson & Holthuis (1955) provide details on the publica-tion dates of the parts containing reptiles. We here are concerned only with pages 1-28 which deal with crocodilians. Pages 1-8 and plates 1-3 were published on December 18, 1839, pages 9-28 on October 11, 1841. We have used these dates, instead of the general-ised date 1844 found in the literature. Pictures.― Müller & Schlegel, 1839: pl. 3, figs 7, 8.

† Crocodilus ossifragus Dubois, 1908

*Crocodilus ossifragus Dubois, 1908: 1269; Massimo & Vos, 2010.

Syntypes: (all material Java, Indonesia) RMNH.Dub 10, 5 teeth, Trinil; RMNH.Dub 11, skull, Trinil; RMNH.Dub 15, mandible, Trinil; RMNH.Dub 16, dentary, Trinil; RMNH.Dub 17, mandible, Trinil; RMNH.Dub 18, dentary, Trinil; RMNH.Dub 19, mandible, Trinil; RMNH.Dub 27, dentary, Trinil; RMNH.Dub 28, maxilla, Trinil; RMNH.Dub 53, angular, Trinil; RMNH.Dub 74, premaxilla, Trinil; RMNH.Dub 1477, 33 teeth, Trinil; RMNH.Dub 2005, tooth, Trinil; RMNH.Dub 2184, 57 teeth Trinil, RMNH.Dub 2707, 22 teeth, Trinil; RMNH.Dub 10111a, dentary, Trinil; RMNH.Dub 10544, mandible, Trinil; RMNH.Dub 1617b, pterygoid and ectopterygoid, Trinil; RMNH.Dub 45, skull, Bogo; RMNH.Dub 47, skull, Bogo; RMNH.Dub 50, premaxilla, Bogo; RMNH.Dub 6975, tooth, Jeruk; RMNH.Dub 33, 3 teeth, Kali Gedeh; RMNH.Dub 88, 2 teeth, Kali Gedeh; RMNH.Dub 12, skull, Kedung Brubus; RMNH.Dub 21, angular, Kedung Brubus; RMNH.Dub 46, maxilla, Kedung Brubus; RMNH.Dub 48, maxilla, Kedung Brubus; RMNH.Dub 2716, 10 teeth, Kedung Brubus; RMNH.Dub 9940a, osteoderm, Kedung Brubus; RMNH.Dub 42, maxilla, Kebon Duren; RMNH.Dub 42, maxilla, Kebon Duren; RMNH.Dub 12942, tooth, Kebon Duren; RMNH.Dub 29, premaxilla, Padas Malang; RMNH.Dub 13, skull, Teguan; RMNH.Dub 24, skull and mandible, Teguan; RMNH.Dub 37a, 6 dentaries, Teguan; RMNH.Dub 2706, skull, Teguan; RMNH.Dub 20, skull, Java; RMNH.Dub 36, tooth, Java; RMNH.Dub 43, maxilla, Java; RMNH.Dub 44, skull, Java; RMNH.Dub 51, skull, Java; RMNH.Dub 55, angular, Java; RMNH.Dub 2710, 4 teeth, Java; RMNH.Dub 8132, mandible, Java; RMNH.Dub 9945c, osteoderm, Java; RMNH.Dub 10318a, osteoderm, Java; RMNH.Dub 10341c, 3 osteoderms; RMNH.Dub 12945, skull, Java; RMNH.Dub


12946, skull, Java; RMNH.Dub 12948, jugal, Java; RMNH.Dub 12950, angular, Java; RMNH.Dub 12951, dentary, Java; RMNH.Dub 12952, premaxilla, Java; RMNH.Dub 1614, dentary, Java. All localities are of late Early Pleistocene or Middle Pleistocene age.

Current name: Crocodylus siamensis Schneider, 1801. Remarks.― Dubois (1908) only gave a short diagnosis of this species. Janensch (1911) provided an extensive description of material of this species obtained by the Ger-man Trinil expedition. Müller (1923) synonymised Crocodylus ossifragus with the extant Crocodylus siamensis Schneider, 1801. A full description of the material in the Dubois collection is given by Massimo & de Vos (2010). Pictures.― Massimo & Vos (2010): figs 6-9, 17S-29S.

Crocodilus (Gavialis) Schlegelii Müller, 1838(figs 12, 13)

*Crocodilus (Gavialis) Schlegelii Müller, 1838: 77, pl. 3; Müller & Schlegel, 1839: pls. 1-3; Müller & Schlegel, 1841: 18.

Tomistoma schlegelii; Lidth de Jeude, 1898: 13; King & Burke, 1989: 15.

Syntypes: RMNH.RENA 3198, 1 egg in alcohol, ‘Borneo’ [probably Lake Lamoeda, 8 days upstream river Doeson, Borneo, Indonesia], leg. S. Müller, 1837; RMNH.RENA 3200, 1 embryo (alc.). Loc.: ‘Borneo’ [probably Lake Lamoeda, 8 days upstream river Doeson, Borneo, Indonesia]. Leg.: S. Müller; RMNH.RENA 7934-5, 2 mounted ex. Loc.: ‘Borneo’ [probably Lake Lamoeda, 8 days upstream river Doeson, Borneo, Indonesia]. Leg.: S. Müller; RMNH.RENA 35444 (Cat. ost. b, 76 cm), 1 skull. Loc.: ‘Borneo’ [probably Lake Lamoeda, 8 days upstream river Doeson, Borneo, Indonesia]. Leg.: S. Müller, 1836; RMNH.RENA 35445 (Cat. ost. c, 70 cm), 1 skull. Loc.: ‘Dans d’un lac d’eau douce à l’intérieur de Bornéo’ [probably Lake Lamoeda, 8 days upstream river Doeson, Borneo, Indonesia]. Leg.: S. Müller, 1836; RMNH.RENA 35449, 1 dried egg. Loc.: none given [probably Lake Lamoeda, 8 days upstream river Doeson, Borneo, Indonesia]. Leg.: S. Müller; RMNH.RENA 39581 (Cat. ost. a, skull 63.5 cm, total 360

Fig. 12. Tomistoma schlegelii, RMNH.RENA 3200, alcohol preserved embryo, syntype of Crocodilus (Gavialus) schlegelii.


cm), 1 skeleton. Loc.: ‘Borneo’ [probably Lake Lamoeda, 8 days upstream river Doeson, Borneo, Indone-sia]. Leg.: S. Müller.

Current name: Tomistoma schlegelii (Müller, 1838). Remarks.― The type locality according to Müller (1838: 85) is ‘binnenlanden van Borneo’ [interior of Borneo]. Müller obtained eggs with embryos from a nest discovered by P.W. Korthals (also a member of the Dutch Natuurkundige Commissie) in Septem-ber 1836, by Lake Lamoeda (Müller, 1838: 36). Of these eggs apparently only two sur-vived till today: RMNH.RENA 3198 (alc.) and RMNH.RENA 35449 (dry). RMNH.RENA 35449 now is damaged (broken and partly repaired in April 2003). This egg was only found recently in the RMNH collection without any data, but it did agree com-pletely in size and shape with the egg depicted by Müller & Schlegel (1841) and is as-sumed to be one of the eggs mentioned in the original description by Müller (1838) and in Müller & Schlegel (1841). King & Burke (1989: 15) only mention RMNH.RENA 3200, 7934, 7935 and the skel-etal material Cat. ost. a, b and c as syntypes. Pictures.― Müller, 1838: 77, pl. 3; Müller & Schlegel, 1839: pls. 1-3.

Gavialidae Adams, 1854† Garialis bengawanicus Dubois, 1908

*Garialis [= Gavialis] bengawanicus Dubois, 1908: 1269.

Syntypes: (all material Java, Indonesia) RMNH.Dub 1, 2 skulls, Trinil; RMNH.Dub 2, dentary, Trinil; RMNH.Dub 5, rostrum, Trinil; RMNH.Dub 7, mandible, Trinil; RMNH.Dub 8, dentary, Trinil; RMNH.Dub 9 + 1617a, skull, Trinil; RMNH.Dub 10, 86 teeth, Trinil; RMNH.Dub 34b, 33 teeth, Trinil; RMNH.Dub 246, 3 jugals and 2 skull fragments; RMNH.Dub 1473c, 162 teeth, Trinil; RMNH.Dub 1477, 2 teeth; RMNH.Dub 1615, 190 teeth, Trinil; RMNH.Dub 1616, skull table and 10 teeth; Trinil; RMNH.Dub 1618,

Fig. 13. Tomistoma schlegelii, RMNH.RENA 3198, alcohol preserved egg, syntype of Crocodilus (Gavialus) schlegelii.


supraoccipital, Trinil; RMNH.Dub 1620, dentary; Trinil; RMNH.Dub 1621a, dentary, Trinil; RMNH.Dub 1623, skull, Trinil; RMNH.Dub 2655, 68 teeth, Trinil; RMNH.Dub 6401, axis, Trinil; RMNH.Dub 33, tooth, Kali Gedeh; RMNH.Dub 37b, mandible, Kedung Brubus; RMNH.Dub 2716, tooth, Kedung Bru-bus; RMNH.Dub 6 + 25, mandible, Kedung Lumbu; RMNH.Dub 23, 2 splenials, Ngandjar; RMNH.Dub 4 + 32 + 2715, mandible, Pitu; RMNH.Dub 37a, dentary, Teguan; RMNH.Dub 1461a, 8 teeth, Teguan or Bogo; RMNH.Dub 1614, mandible, Java; RMNH.Dub 1621b, skull fragment, Java; RMNH.Dub 12947, skull fragment, Java; RMNH.Dub 15583, osteoderm, Java, RMNH.Dub 15584, 3 osteoderms, Java. All localities are of late Early Pleistocene or Middle Pleistocene age.

Current name: † Gavialis bengawanicus Dubois, 1908. Remarks.― Dubois (1908) clearly wrote Garialis with a ‘r’ instead of a ‘v’. This is clear from comparison with other italicized names with a ‘r’ or a ‘v’. However, Dubois (1908) compared his Garialis bengawanicus with ‘G. gangeticus’, a species for which only the generic name Gavialis ever has been used. We therefore infer that the use of the ‘r’ instead of a ‘v’ was inadvertent, and due to misinterpretation of a handwritten manu-script by the printer. Therefore there are no nomenclatural consequences. Dubois (1908) only gave a short diagnosis of this species. Janensch (1911) provided an extensive description of the material of this species obtained by the German Trinil expedition. A full description of the material in the Dubois collection is given by Mas-simo & de Vos (2010). Pictures.― Massimo & De Vos (2010): figs 1-5, 2S-15S.

Specimens in the RMNH collections indicated as types, which are not types, or types of other names

‘Chelodina rottiensis nov. sp.’ RMNH.RENA 10187, RMNH.RENA 4349 from Rotti, Indonesia, collected by H.C.F. ten Kate. As mentioned under Chelodina mccordi this is a label name coined by L.D. Brongersma that was never published. However, RMNH.RENA 10187 is a paratype of Chelodina mccordi Rhodin.

‘Cistudo gastrotaenia Bleeker’ RMNH.RENA 4063 (alc.) contains two hatchlings of Cuora amboinensis (Daudin, 1801) and on the label has the handwritten indication ‘Cistudo gastrotaenia, type, Dr. Bleeker, Arch. Ind.’ We have not been able to find any published record of this name, and have to assume that it is a manuscript or label name of Dr. P. Bleeker that was never published and nomenclatorially does not exist.

Clemmys Sigriz Michahelles, 1829 RMNH 3327 erroneously has been indicated as type of this name. For further com-ments see under Emys vulgaris Gray, 1831.

Emydura Adolf Friedr[ich] spec. nov. Kopstein RMNH.RENA 20700, Chelodina cf. branderhorsti (Ouwens, 1914) from the Jakati River, Bintoemi Bay, SW Dutch New Guinea, October 3, 1923 collected by Dr. F. Kop-stein, bears a partly legible label showing this name. We have not been able to locate this name in literature and assume this is a label and/or manuscript name coined by F. Kopstein after his collecting trip to the Jakati River.


Notochelys platynota (Gray, 1834) RMNH.RENA 3348 on a rather recently handwritten label is incorrectly is indicated as the holotype of this species, but this specimen is a syntype of Cyclemys giebelii only (see above). This, however, is due to an administrative mistake of a technician writing a wrong text (Fig. 1). This misunderstanding has never entered the literature.

Conclusions and summary

We could locate 31 primary types, viz. five holotypes, five lectotypes and 21 syn-types of ten names (not counting all the fossilized segments of two crocodilian taxa), belonging to in total 19 species. Holotypes: 5 (Chelonura Temminckii, RMNH.RENA 6166; †Hardella isoclina RMNH Dub. 2722; Testudo Forstenii RMNH.RENA 3811; Testudo Strauchi RMNH.RENA 6011; Testudo indica Vosmaeri RMNH.RENA 6001) Lectotypes: 5 (Emys subtrijuga RMNH.RENA 6082; Testudo dussumieri RMNH.RENA 3231; Testudo emys RMNH.RENA 3808; Emys macrocephala RMNH.RENA 6164; Crocodylus biporcatus raninus RMNH.RENA 3219). Syntypes: 21 (Cyclemys giebelii RMNH.RENA 3348; Emys borneoensis RMNH.RENA 3296, 6210; Emys vulgaris japonica RMNH.RENA 3331, 3332, 3333 (2 ex.), 3334, 6142; Emys vulgaris picta RMNH.RENA 3330 (2 ex.); Trionys stellatus var. japonicus RMNH.RENA 3259, 3264; †Crocodilus ossifragus RMNH Dub. a large collection of fossilized re-mains, no complete specimen; Tomistoma Schlegelii RMNH.RENA 3198, 3200, 7934, 7935, 35444, 35445, 35449, 39581; †Garialis bengawanicus RMNH Dub. a large collection of fos-silized remains, no complete specimen, but one incomplete skull. The following 26 secondary types were located, viz., 12 paratypes of six names and 14 paralectotypes of six names. Paratypes: 12 (Emys orbicularis occidentalis RMNH.RENA 11371 (2 ex.), 15003; Cyclemys enigmatica RMNH.RENA 3838, 6066, 6068, 27828; Cyclemys pulchristriata RMNH.RENA 4751; Geoemyda spengleri sinensis RMNH.RENA 5887; Batrachemys heliostemma RMNH.RENA 31998, 31999; Chelodina mccordi RMNH.RENA 10187). Paralectotypes: 14 (Emys dentata RMNH.RENA 6062, 6063, 6067, 40474; *Emys subtrijuga RMNH.RENA 6084, 6085; Emys vulgaris RMNH.RENA 3327: *Testudo emys RMNH.RENA 6005, 6030, 17967; Emys Amazonica RMNH.RENA 3294; *Crocodylus biporcatus raninus RMNH.RENA 7939+21695 (skin and skull + lower jaw of same specimen), 37489, 37493). Of names indicated with * the lectotype also is present in the RMNH collection. A total of 31 primary and 26 secondary type specimens was located for 30 nominal names, of which 17 (15 recent and two fossil taxa) still are recognised as full species today. We think that this inventory is complete and that among the RMNH material there are no more types hiding. The type localities of the RMNH type material are spread over the world: Europe (1 specimen, 1 taxon), N. America (1 specimen, 1 taxon), S. America (4 specimens, 3 taxa), Africa (4 specimens, 2 taxa), Aldabra (1 specimen, 1 taxon), Mascarenes (1 specimen, 1 taxon), mainland SE Asia (2 specimens, 2 taxa), Indonesia (33 specimens of 13 nominal taxa, not counting the material of 2 extinct crocodilians), Japan (10 specimens, 3 taxa), although there is a clear majority, both in number of specimens and number of taxa from Indonesia. The fact that the RMNH collections contain the type series of two recent crocodilian taxa (8.3% of existing recent taxa) is noteworthy.


Acknowledgements

A first skeleton draft of this type list was prepared by Dr. L.W. van den Hoek Os-tende (RMNH) under the supervision of Dr. M.S. Hoogmoed in the period 1995/96, during work on the Delta Plan for the Preservation of Cultural Heritage, a project fi-nanced by the then Ministerie van Welzijn, Volksgezondheid en Cultuur (= Ministry of Welfare, Public Health and Culture). This project provided Dutch National Museums, among which the RMNH, ‘with extra funding above their normal operating budgets to reduce arrears in conservation and registration. It provided a unique possibility to reg-ister type-material and thus fulfil recommendation 72G (4) of the International Code of Zoological Nomenclature, viz., that ‘every institution in which name-bearing types are deposited should publish lists of name-bearing types in its possession or custody’. The registration of types and specimens of extinct or endangered species in the vertebrate collections [was] one of the many projects in the National Museum of Natural History financed by the ‘Delta Plan’’(Hoek Ostende et al., 1997). Prof. Dr. U. Fritz provided use-ful suggestions about the status of old material of some European and Southeast Asian species of Mauremys and Cyclemys. R. de Ruiter, collection manager of the herpetologi-cal collection of the National Museum of Natural History, Leiden, The Netherlands, provided information about recent herpetological material in the RMNH collection. We want to thank Dr. J. de Vos and R. van Zelst of the department of Geology of the National Museum of Natural History for the help and the information they provided about the fossil Dubois type material in the RMNH collection. The present work has been partly financed by a grant from the SenterNovem Type Project (project no. 1280).

References

Boeseman, M., 1970. The vicissitudes and dispersal of Albertus Seba’s zoological specimens.― Zoolo-gische Mededelingen, Leiden 44: 177-206.

Bour, R., 1984. L’identité de Testudo gigantea Schweigger, 1812 (Reptilia, Chelonii).― Bulletin du Muséum National d’Histoire Naturelle, Paris 4 Sér., 6, section A(1):159-175.

Bour, R. 1987. Type-specimen of the Alligator Snapper, Macroclemys temminckii (Harlan, 1835).― Journal of Herpetology 21: 340-343.

Bour, R., 2006. Identity of Testudo gigantea Schweigger, 1812 and rediscovery of the type specimen.― Emys 13: 12-23.

Bour, R. & P.C.H. Pritchard, 2009. Comments on the proposed conservation of usage of Testudo gigantea Schweigger, 1812 (currently Geochelone (Aldabrachelys) gigantea) (Reptilia, Testudines) (Case 3463) 1.― Bulletin Zoological Nomenclature 66: 169-174.

Bour, R, J.B. Iverson & P.C.H. Pritchard, 2010. Comments on the proposed conservation of usage of Testudo gigantea Schweigger, 1812 (currently Geochelone (Aldabrachelys) gigantea) (Reptilia, Testu-dines)(Case 3463) (2). Jean-Jacques Dussumier and Aldabra.― Bulletin Zoological Nomenclature 67: 72-73.

Brophy, T.R., 2004. Geographic variation and systematics in the South-east Asian turtles of the genus Malayemys (Testudines: Bataguridae).― Hamadryad 29: 63-79.

Cheke, A.S., 2009. Comments on the proposed conservation of usage of Testudo gigantea Schweigger, 1812 (currently Geochelone (Aldabrachelys) gigantea) (Reptilia, Testudines) (Case 3463-2).― Bulletin Zoological Nomenclature 66: 174-176.

Cheke, A., 2010. Comments on the proposed conservation of usage of Testudo gigantea Schweigger, 1812 (currently Geochelone (Aldabrachelys) gigantea) (Reptilia, Testudines)(Case 3463-6).― Bulletin Zoo-logical Nomenclature 67: 79-81.


Dubois, E., 1908. Das Geologische Alter der Kendeng- oder Trinil-Fauna.― Tijdschrift van het Koninklijk Nederlandsch Aardrijkskundig Genootschap, 2nd series 35(6): 1235-1270.

Dubois, A., A. Ohler & E.-R. Brygoo, 2010. Comments on the proposed conservation of usage of Testudo gigantea Schweigger, 1812 (currently Geochelone (Aldabrachelys) gigantea) (Reptilia, Testudines)(Case 3463-8).― Bulletin Zoological Nomenclature 67(1): 82-89.

Duméril, A.M.C. & A. Duméril, 1851. Catalogue méthodique de la collection des Reptiles du Muséum d’Histoire Naturelle de Paris: i-iv, 1-224.― Gide et Baudry, Paris.

Fan, T.H., 1931. Preliminary report of Reptiles from Yaoshan, Kwangsi, China.― Bulletin of the Depart-ment of Biological Collections and Sciences Sun Yatsen Univ. Canton 11: 1-154.

Fitzinger, L.I., 1826. Neue Classification der Reptilien nach iheren naturlichen Verwandtschaften. Nebst einer Verwandtschaftstafel und einem Verzeichnisse der Reptilien der Reptilien-Sammlung des K.K. Zoologischen Museums zu Wien: 1-66.― Vienna.

Fitzinger, L.I., 1835. Entwurf einer systematischen Anordnung der Schildkröten nach den Grund-sätzen der natürlichen Methode.― Annalen des Wiener Museums der Naturgeschichte 1(6): 103-128.

Fransen, C.H.J.M., L.B. Holthuis & J.P.H.M.Adema, 1997. Type-catalogue of the Decapod Crustacea in the collection of the Nationaal Natuurhistorische Museum, with appendices of pre-1900 collectors and material.― Zoologische Verhandelingen 311: i-xvi, 1-344.

Franzen, M. & F. Glaw, 2007. Type catalogue of reptiles in the Zoologische Staatssammlung München. Spixiana 30: 201-274.

Frazier, J., 2006. A neotype for the Aldabra tortoise, Testudo gigantea Schweigger, 1812.― Herpetological Review 37: 275-280.

Frazier, J., 2009. Case 3463. Testudo gigantea Schweigger, 1812 (currently Geochelone (Aldabrachelys) gigantea: Reptilia, Testudines): proposed conservation of usage of the specific name by maintenance of a designated neotype, and suppression of Testudo dussumieri Gray, 1831 (currently Dipsochelys dussumieri).― Bulletin of Zoological Nomenclature 66: 34-50.

Fritz, U. 1993. Zur innerartlichen Variabilität von Emys orbicularis (Linnaeus, 1758). 3. Zwei neue Unter-arten von der Iberischen Halbinsel und aus Nordafrika, Emys orbicularis fritzjuergenobsti subsp. nov. und E. o. occidentalis subsp. nov.― Zoologische Abhandlungen Museum für Tierkunde Dresden 47: 131-155.

Fritz, U., M. Gaulke & E. Lehr, 1997. Revision der südostasiatischen Dornschildkröten-Gattung Cyclemys Bell, 1834, mit Beschreibung einer neuen Art.― Salamandra 33: 183-212.

Fritz, U., S. Gong, M. Auer, G. Kuchling, N. Schneeweiß, A.K. Hundsdörfer, 2010. The world´s eco-nomically most important chelonians represent a diverse species complex (testudines: Trionychi-dae: Pelodiscus).― Organisms Diversity & Evolution: DOI 10.1007/s13127-010-0007-1.

Fritz, U., D. Guicking, M. Auer, R.S. Sommer, M. Wink & A.K. Hundsdörfer, 2008. Diversity of the Southeast Asian leaf turtle genus Cyclemys: how many leaves on its tree of life? – Zoologia Scripta 37: 367-390.

Fritz, U. & P. Havaš, 2007. Checklist of Chelonians of the world.― Vertebrate Zoology 57: 149-368. [also as CITES document: 1-230, available at: http://www.cites.org/common/com/NC/tax_ref/Cheloni-ans_Checklist_2006.pdf].

Fritz, U. & T. Wischuf, 1997. Zur Systematik westasiatisch-südosteuropäischer Bachschildkröten (Gat-tung Mauremys) (Reptilia: Testudines: Bataguridae).― Zoologische Abhandlungen, Staatliches Mu-seum für Tierkunde, Dresden 49: 223-260.

Frost, D.R., 2009. Amphibian species of the world: an online reference. Version 5.3. American Museum of Natural History, New York, USA. [electronic database accessible at http://research.amnh.org/herpetology/amphibia/index.php., accessed 12 February 2009].

Gassó Miracle, M.E., L.W. van den Hoek Ostende & J.W. Arntzen, 2007. Type specimens of amphibians in the National Museum of Natural History, Leiden, The Netherlands.― Zootaxa 1482: 25-68.

Gerlach, J., 2004. Giant tortoises of the Indian Ocean. The genus Dipsochelys inhabiting the Seychelles Islands and the extinct giants of Madagascar and the Mascarenes: 1-207.― Edition Chimaira, Frankfurt am Main.


Giebel, C., 1866. Die Schildkröten der Insel Banka.― Zeitschrift für die Gesammten Naturwissenschaft: 11-21.

Gijzen, A., 1938. ‘s Rijks Museum van Natuurlijke Historie, 1820-1915: i-xii, 1-335.― Dissertation. Am-sterdam, W.L. en J. Brussen.

Gray, J.E., 1831a. A synopsis of the species of the class Reptilia. In: E. Griffith & E. Pidgeon: The animal kingdom arranged in conformity with its organization, by the Baron Cuvier, with additional de-scriptions of all the species hitherto named, and of many not before noticed. Vol. 9. The class Rep-tilia arranged by the Baron Cuvier, with specific descriptions: 1-110 (second set of pages).― Whit-taker, Treacher, and Co., London

Gray, J.E., 1831b. Synopsis Reptilium or short descriptions of the species of reptiles. Part I. Cataphracta. Tortoises, Crocodiles and Enaliosaurians: i-viii, 1-88.― London.

Gray, J.E., 1844. Catalogue of the Tortoises, Crocodiles, and Amphisbaenians, in the collection of the British Museum: i-viii, 1-80.― Edward Newman, London.

Gray, J.E., 1873. Hand-list of the specimens of shield reptiles in the British Museum: i-iv, 1-124.― Trus-tees of the British Museum, London.

Grünewald, F., 2009a. Museumcollecties.RMNH 31998 & 31999 (Mesoclemmys heliostemma) McCord, Joseph-Ouni & Lamar, 2001.― Trionyx 7: 16-21.

Grünewald, F., 2009b. Museumcollecties.RMNH 3231 (Dipsochelys dussumieri) Gray, 1831.― Trionyx 7: 142. Günther, A., 1877. The gigantic tortoises (living and extinct) in the collection of the British Museum:

1-96, pls. 1-54. London.Harlan, R., 1835. Genera of North American Reptilia, and a synopsis of the species.― Medical and

Physical Researches or original memoirs in medicine, surgery, physiology, geology, zoology, and comparative anatomy 1835: 84-163.

Hoek Ostende, L.W. van den, R.W.R.J. Dekker & G.O Keijl, 1997. Type-specimens of birds in the Na-tional Museum of Natural History, Leiden.― NNM Technical Bulletin 1: 1-248.

Hoek Ostende, L.[W.] van den, C. Smeenk, R.[W.R.J.] Dekker, M.[S.] Hoogmoed, M. van Oijen & R. Pur-cell (pictures), 1999. Swift as a shadow: 1-159.― Houghton Mifflin Company, Boston and New York.

Holbrook, J.E., 1842. North America Herpetology or a description of the Reptiles inhabiting the United States. Vol. I (second edition): i-xv, 17-152.― Dobson: Philadelphia.

Holthuis, L.B., 1995. 1820-1958 Rijksmuseum van Natuurlijke Historie: 1-172.― Leiden, Nationaal Na-tuurhistorisch Museum.

Holthuis, L.B., & T. Sakai, 1970. Ph.F. von Siebold and Fauna Japonica. A history of early Japanese Zool-ogy: 1-323.― Academic Press of Japan

Hoogmoed, M.S., 1982. Nomenclatural problems relating to Atractus trilineatus Wagler, 1828.― Zoolo-gische Mededelingen Leiden 56: 131-138.

Hoogmoed, M.S., 1999. Rodriguez Giant Tortoise. In: Hoek Ostende, L.[W.] van den, C. Smeenk, R.[W.R.J.] Dekker, M.[S.] Hoogmoed, M. van Oijen & R. Purcell (pictures): Swift as a shadow: 57.― Houghton Mifflin Company, Boston and New York.

Hoogmoed, M.S. & C.R. Crumly, 1984. Land tortoise types in the Rijksmuseum van Natuurlijke Historie with comments on nomeclature and systematics (Reptilia: Testudines: Testudinidae).― Zoolo-gische Mededelingen Leiden 58 (15): 241-259.

Hoogmoed, M.S. & U. Gruber, 1983. Spix and Wagler type specimens of reptiles and amphibians in the Natural History Musea in Munich (Germany) and Leiden (The Netherlands).― Spixiana 9 suppl.: 319-415.

Hubrecht, A.A.W., 1881. On certain tortoises in the collections of the Leiden Museum.― Notes from the Leyden Museum 3: 41-50.

Husson, A.M. & L.B. Holthuis, 1955. The dates of publication of ‘Verhandelingen over de Natuurlijke Geschiedenis der Nederlandsche Overzeesche bezittingen’ edited by C.J. Temminck.― Zoologische Mededelingen 34: 17-24.

International Commission on Zoological Nomenclature, 1999. International Code of Zoological Nomen-clature (Fourth Edition): i-xxix, 1-306.

Jaeckel, O., 1911. Die fossilen Schildkrötenreste von Trinil. In: Selenka, M. & M. Blankenhorn: Die Pithe-canthropus-Schichten auf Java. Geologische und Paläontologische Ergebnisse der Trinil Expedition


(1907 und 1908). Ausgeführt mit Unterstützung der Akademischen Jubiläumsstiftung der Stadt Berlin und der Königlich Bayerischen Akademie der Wissenschaften: 75-81.― Verlag Wilhelm Engelmann, Leipzig.

Janensch, W., 1911. Die Reptilienreste (exkl. Schildkröten). In: Selenka, M. & M. Blankenhorn: Die Pithe-canthropus-Schichten auf Java. Geologische und Paläontologische Ergebnisse der Trinil Expedition (1907 und 1908). Ausgeführt mit Unterstützung der Akademischen Jubiläumsstiftung der Stadt Berlin und der Königlich Bayerischen Akademie der Wissenschaften: 61-74.― Verlag Wilhelm Engelmann, Leipzig.

Kate, H.F.C. ten, 1894. Verslag eener reis in de Timorgroep en Polynesie. IV. Roti.-Savoe.― Tijdschrift van het Koninklijk Nederlandsch Aardrijkskundig Genootschap (2) 11: 659-700.

King, F.W. & R.L. Burke (eds.), 1989. Crocodilian, Tuatara, and Turtle Species of the World: A Taxo-nomic and Geographic Reference: i-xxii, 1- 216.― Association of Systematics Collections, Washing-ton, D.C. [available on line: http://www.flmnh.ufl.edu/natsci/herpetology/turtcroclist/, accessed 7 April 1997].

Lidth de Jeude, T.W. van, 1893. Note XLIII. On a new species of the genus Testudo.― Notes from the Leyden Museum 15: 312-313.

Lidth de Jeude, T.W. van, 1895. Reptiles from Timor and the neighbouring islands.― Notes from the Leyden Museum 16: 119-127.

Lidth de Jeude, T.W. van, 1896. Note XXVIII. On Testudo emys Schleg. & Mull. and its affinities.― Notes from the Leyden Museum 17: 197-204, pls. 5-6.

Lidth de Jeude, T.W. van, 1898. Museum d’Histoire Naturelle des Pays Bas. Tome X2. Seconde Partie. Catalogue Ostéologique des poissons, reptiles et amphibiens. Reptiles: 1-52.

Massimo, M. & J. de Vos, 2010. A revision of Dubois crocodylians, Gavialis bengawanicus and Crocodylus ossifragus, from the Pleistocene Homo erectus beds of Java.― Journal of Vertebrate Paleontology, 30: 427-441 + Supplementary Data 1 (30 pp.) and 2 (1 p.).

Matyot, P., 2009. Comments on the proposed conservation of usage of Testudo gigantea Schweigger, 1812 (currently Geochelone (Aldabrachelys) gigantea) (Reptilia, Testudines)(Case 3463-1).― Bulletin Zoo-logical Nomenclature 66: 352-354.

McCord, W.P., M. Joseph-Ouni & W.W. Lamar, 2001. A Taxonomic Reevaluation of Phrynops (Testudines: Chelidae) with the description of two new genera and a new species of Batrachemys.― Revista de Biologia Tropical 49: 715-764.

Merrem, B., 1820. Versuch eines Systems der Amphibien. 1-191.― Marburg Michahelles, C.W., 1829. Commentatio de speciebus aut rarioribus, aut novis cheloniorum Europam

meridionalem inhabitantibus.― Isis (Oken), Leipzig (1829): 1295-1300.Müller, L., 1923. Crocodylus siamensis Schneid. und Crocodylus ossifragus Dubois.― Palaeontologica Hun-

garica 1: 109-122.Müller, S., 1838. Waarnemingen over de Indische krokodillen en beschrijving van eene nieuwe soort.―

Tijdschrift voor Natuurlijke Geschiedenis en Physiologie 5: 61-87.Müller, S. & H. Schlegel, 1839. Over de krokodillen van de Indischen Archipel. In: Temminck, C.J. (ed.),

Verhandelingen over de Natuurlijke Geschiedenis der Nederlandsche Overzeesche bezittingen. 1839-1844. Zoologie. Vol. 2: 1-8, pls. 1-3.― Leiden.

Müller, S. & H. Schlegel, 1841. Over de krokodillen van de Indischen Archipel. In: Temminck, C.J. (ed.), Verhandelingen over de Natuurlijke Geschiedenis der Nederlandsche Overzeesche bezittingen. 1839-1844. Zoologie. Vol. 2: 9-28.― Leiden.

Opinion 660. ICZN, 1963. Suppression under the plenary powers of seven specific names of turtles (Reptilia, Testudines).― The Bulletin of Zoological Nomenclature 20(3):187-190.

Praschag, P., A.K. Hundsdörfer & U. Fritz, 2007. Phylogeny and taxonomy of endangered South and South-east Asian freshwater turtles elucidated by mtDNA sequence variation (Testudines: Geoe-mydidae: Batagur, Callagur, Hardella, Kachuga, Pangshura).― Zoologica Scripta 36: 429-442.

Pritchard, P.C.H., 1986. A reinterpretation of Testudo gigantea Schweigger, 1812.― Journal of Herpetolo-gy 20: 523-534

Pritchard, P.C.H., 1989. The alligator snapping turtle. Biology and conservation: i-xi, 1-104.― Milwau-kee, WI: Milwaukee Public Museum.


Rhodin, A.G.J., 1994. Chelid turtles of the Australasian archipelago: II. A new species of Chelodina from Roti Island, Indonesia.― Breviora 498: 1-31.

Rooij, N. de, 1915. The reptiles of the Indo-Australian Archipelago. I. Lacertilia, Chelonia, Emydosauria: i-xiv, 1-384.― E.J. Brill, Leiden.

Ross, C.A., 1990. Crocodylus raninus S. Muller and Schlegel, a valid species of crocodile (Reptilia: Croco-dylidae) from Borneo.― Proceedings of the Biological Society of Washington 103 (4): 955-961.

Ross, C.A., 1992. Designation of a lectotype for Crocodylus raninus S. Müller and Schlegel (Reptilia: Croc-odylidae), the Borneo crocodile.― Proceedings of the Biological Society of Washington 105: 400-402.

Schlegel, H., 1837-1844. Abbildungen neuer oder unvollständig bekannter Amphibien, nach der Natur oder dem Leben entworfen: 1-141.― Arnz and Comp., Düsseldorf.

Schlegel, H. & S. Müller, 1840. Over de schildpadden van den Indische Archipel, en beschrijving eener nieuwe soort van Sumatra. In: Temminck, C.J. (ed.), Verhandelingen over de Natuurlijke Ge-schiedenis der Nederlandsche Overzeesche bezittingen. 1839-1844. Zoologie. Vol. 2: pl.4. Leiden.

Schlegel, H. & S. Müller, 1845. Over de schildpadden van den Indische Archipel, en beschrijving eener nieuwe soort van Sumatra. In: Temminck, C.J. (ed.), Verhandelingen over de Natuurlijke Ge-schiedenis der Nederlandsche Overzeesche bezittingen. 1839-1844. Zoologie. Vol. 2: 29-36. Leiden.

Schoepff, J.D., 1792. Historia Testudinum: i-xii, 1-136, pls. 1-31. Erlangen.Schoepff, J.D., 1801. Naturgeschichte der Schildkröten mit Abbildungen: 1-160, pls.1-32. Erlangen.Sherborn, C.D. & F.A. Jentink, 1895. On the dates of the parts of Siebold’s ‘Fauna Japonica’ and Giebel’s

‘Allgemeine Zoologie’ (first edition).― Proceedings of the Zooogical. Society of London: 149-150.Spix, J.B. von, 1824. Animalia nova sive species novae Testudinum et Ranarum, quas in itinere per Bra-

siliam annis MDCCCXVII-MDCCCXX: 1-53.― München.Stejneger, L., 1907. The herpetology of Japan and adjacent territory.― Bulletin of the U.S. National Mu-

seum, 58: i-xx, 1-577.Strauch, A., 1865. Die Vertheilung der Schildkröten uber den Erdball.― Mémoire de l’Académie Impé-

riale des Sciences de St.-Pétersbourg, 7 serie, Tome 8 (13): 12-207.Strauch, A., 1866. Synopsis der gegenwärtig lebenden Crocodiliden.― Mémoire de l’Académie Impé-

riale des Sciences de St.-Pétersbourg, 7 serie, Tome 10 (13): 1-120.Suckow, G.A., 1798. Anfangsgründe der theoretischen und angewandten Naturgeschichte der Thiere

Dritter Theil, von den Amphibien: 1-298.― Weissman, Leipzig.Temminck, C.J.& H. Schlegel. 1834. Reptilia. In: P.F. von Siebold, Fauna Japonica sive Descriptio animal-

ium, quae in itinere per Japonianum, jussu et auspiciis superiorum, qui summum in India Batava Imperium tenent, suscepto, annis 1823-1830 colleget, notis observationibus et adumbrationibus il-lustratis. Vol. III: i-xxii, 1-144, pls. 1-27.― J.G. Lalau, Leiden.

Vaillant, L., 1893. Les tortues éteintes de l’ile Rodriguez d’après les pièces conservées dans les galeries du muséum.― Volume commémoratif du Centenaire de la fondation du Muséum d’Histoire Na-turelle: 1-37.

Vervoort, W., 1981. Verslag van de directeur over het jaar 1980. Rijksmuseum van Natuurlijke Historie te Leiden. Nederlandse Rijksmusea 102: 195-250.

Wermuth, H. & R. Mertens, 1961. Schildkröten. Krokodile. Brückenechsen: i-xxiv, 1-422.― Gustav Fischer Verlag, Jena.

Williams, E.E., 1957. Hardella isoclina Dubois redescribed.― Zoologische Mededelingen Leiden 35: 235-240.

Received: 11.xi.2009Accepted: 31.v.2010Edited: L.P. van Ofwegen



Alphabetical list of the current names for the type specimens in the RMNH, with their original names (if different) between brackets

Batagur borneoensis (Schlegel & Müller, 1845) [Emys borneoensis Schlegel & Müller, 1845] Batrachemys heliostemma McCord, Joseph-Oudi & Lamar, 2001 Chelodina mccordi Rhodin, 1994 Crocodylus porosus Schneider, 1801 [Crocodilus biporcatus raninus Müller & Schlegel, 1841 part] Crocodylus raninus Müller & Schlegel, 1841 [Crocodilus biporcatus raninus Müller & Schlegel, 1841 part] Crocodylus siamensis Schneider, 1801 [Crocodilus biporcatus raninus Müller & Schlegel, 1841 part; †Crocodylus ossifragus Dubois, 1908] Cyclemys dentata (Gray, 1831) [Emys dentata Gray, 1831 part] Cyclemys enigmatica Fritz, Guicking, Auer, Sommer, Wink & Hundsdörfer, 2008 [Emys dentata Gray, 1831 part] Cyclemys pulchristriata Fritz, Gaulke & Lehr, 1997 Cylindraspis vosmaeri (Suchow, 1798) [Testudo indica Vosmaeri Suchov, 1798] Dipsochelys dussumieri (Gray, 1831) [Testudo dussumieri Gray, 1831] Emys orbicularis occidentalis Fritz, 1993 †Gavialis bengawanicus Dubois, 1908 [†Garialis bengawanicus Dubois, 1908] Geoemyda spengleri (Gmelin, 1789) [Geoemyda spengleri sinensis Fan, 1931] Indotestudo forstenii (Schlegel & Müller, 1845) [Testudo Forstenii Schlegel & Müller, 1845] Macrochelys temminckii (Troost in Harlan, 1835) [Chelonura Temminckii Troost in Har-lan, 1835 Malayemys subtrijuga (Schlegel & Muller, 1845 [Emys subtrijuga Schlegel & Müller, 1845] Manouria emys emys (Schlegel & Müller, 1840) [ Testudo emys Schlegel & Müller, 1840] †Mauremys? isoclina (Dubois, 1908) [†Hardella isoclina Dubois, 1908] Mauremys japonica (Temminck & Schlegel, 1834) [Emys vulgaris japonica Temminck & Schlegel, 1834] Mauremys reevesii (Gray, 1831) [Emys vulgaris picta Schlegel, 1844] Mauremys rivulata (Valenciennes, 1833) [Emys vulgaris Gray, 1831] Pelodiscus sinensis (Wiegmann, 1834) [Trionyx stellatus var. Japonicus Temminck & Schlegel, 1834]. Note that only recently it has been realized that P. sinensis is a species complex, and thus the identification is provisional, pending further study (Fritz et al., 2010). Peltocephalus dumerilianus (Schweigger, 1812) [Emys macrocephala Spix, 1824] Podocnemis expansa (Schweigger, 1812) [Emys Amazonica Spix, 1824] Psammobates geometricus (Linnaeus, 1758) [Testudo Strauchii Lidth de Jeude, 1893] Tomistoma schlegelii (Müller, 1838) [Crocodilus (Gavialis) Schlegelii Müller, 1838]


Alphabetical list of original names of RMNH types with their current identification in bold

Batrachemys heliostemma McCord, Joseph-Ouni & Lamar, 2001 = Mesoclemmys heliostemma (McCord, Joseph-Ouni & Lamar, 2001) Chelodina mccordi Rhodin, 1994 Chelonura Temminckii Troost in Harlan, 1835 = Macrochelys temminckii (Troost in Har-lan, 1835) Crocodilus biporcatus raninus Müller & Schlegel, 1841 = Crocodylus porosus Schneider, 1801 (part), Crocodylus raninus Müller & Schlegel, 1841 (part), Crocodylus siamensis Schnei-der, 1801 (part) †Crocodilus ossifragus Dubois, 1908 = Crocodylus siamensis Schneider, 1801 Crocodilus (Gavialis) Schlegelii Müller, 1838 = Tomistoma schlegelii (Müller, 1838) Cyclemys enigmatica Fritz, Guicking, Auer, Sommer, Wink & Hundsdörfer, 2008 Cyclemys giebelii Hubrecht, 1881 = Notochelys platynota (Gray, 1834) Cyclemys pulchristriata Fritz, Gaulke & Lehr, 1997 Emys Amazonica Spix, 1824 = Podocnemis expansa (Schweigger, 1812) Emys borneoensis Schlegel & Müller, 1841 = Batagur borneoensis (Schlegel & Müller, 1841) Emys dentata Gray, 1831 (part) = Cyclemys dentata (Gray, 1831 (part), Cyclemys enigmatica Fritz, Guicking, Auer, Sommer, Wink & Hundsdörfer, 2008 (part) Emys macrocephala Spix, 1624 = Peltocephalus dumerilianus (Schweigger, 1812) Emys orbicularis occidentalis Fritz, 1993 Emys subtrijuga Schlegel & Müller, 1845 = Malayemys subtrijuga (Schlegel & Müller, 1845)] Emys vulgaris Gray, 1831 = Mauremys rivulata (Valenciennes, 1833) Emys vulgaris japonica Temminck & Schlegel, 1834 = Mauremys japonica (Temminck & Schlegel, 1834) Emys vulgaris picta Schlegel, 1844 = Mauremys reevesii (Gray, 1831) †Garialis bengawanicus Dubois, 1908 = †Gavialis bengawanicus Dubois, 1908 Geoemyda spengleri sinensis Fan, 1931 = Geoemyda spengleri (Gmelin, 1789) †Hardella isoclina Dubois, 1908 = †Mauremys? isoclina (Dubois, 1908) Testudo dussumieri Gray, 1831 = Dipsochelys dussumieri (Gray, 1831) Testudo emys Schlegel & Müller, 1840 = Manouria emys emys (Schlegel & Müller, 1840) Testudo Forstenii Schlegel & Müller, 1845 = Indotestudo forstenii (Schlegel & Müller, 1845) Testudo Strauchii Lidth de Jeude, 1893 = Psammobates geometricus (Linnaeus, 1758) Testudo indica Vosmaeri Suckow, 1798 = Cylindraspis vosmaeri (Suckow, 1798) Trionyx stellatus var. japonicus Temminck & Schlegel, 1834 = Pelodiscus sinensis (Wieg-mann, 1834. Note that only recently it has been realized that P. sinensis is a species com-plex, and thus the identification is provisional, pending further study (Fritz et al., 2010).


Table 1. Summary of nominal and present names, type status and number of type specimens of Testudines and Crocodylia in the RMNH collections.

Number in Original name Author Year Present name Holotype Lectotype Syntypes Para(lecto)types RMNH.RENATESTUDINES Chelydridae Chelonura Temminckii Troost in Harlan 1835 Macrochelys temminckii RMNH.RENA 6166 1Emys orbicularis occidentalis Fritz 1993 Emys orbicularis occidentalis RMNH.RENA 11371, 15003 3Geoemydidae Cyclemys enigmatica Fritz et al. 2008 Cyclemys enigmatica RMNH.RENA 3838, 6066, 4 6068, 27828 Cyclemys giebelii Hubrecht 1881 Notochelys platynota RMNH.RENA 3348 1Cyclemys pulchristriata Fritz et al. 1997 Cyclemys pulchristriata RMNH.RENA 4751 1Emys borneoensis Schlegel & Müller 1845 Batagur borneoensis RMNH.RENA 3296, 6210 2Emys dentata Gray 1831 Cyclemys dentata RMNH.RENA 6063, 6067, 40474 3Emys dentata Gray 1831 Cyclemys enigmatica RMNH.RENA 6062 1Emys subtrijuga Schlegel & Müller 1845 Malayemys subtrijuga RMNH.RENA 6082 RMNH.RENA 6084, 6085 3Emys vulgaris Gray 1831 Mauremys rivulata RMNH.RENA 3327 1Emys vulgaris japonica Temminck & Schlegel 1834 Mauremys japonica RMNH.RENA 3331-3334, 6

6142Emys vulgaris picta Schlegel 1844 Mauremys reevesii RMNH.RENA 3330 A-B, 2

MNHNP 1954 Geoemyda spengleri sinensis Fan 1931 Geoemyda spengleri RMNH.RENA 5889 1† Hardella isoclina Dubois 1908 † Mauremys? isoclina RMNH Dub. 2722 1Testudinidae Testudo dussumieri Gray 1831 Dipsochelys dussumieri RMNH.RENA 3231 1Testudo emys Schlegel & Müller 1840 Manouria emys emys RMNH.RENA 3808 RMNH.RENA 6005, 6030, 17967: 4 MNHNP 9422 Testudo Forstenii Schlegel & Müller 1845 Indotestudo forstenii RMNH.RENA 3811 1Testudo Strauchi Lidth de Jeude 1893 Psammobates geometricus RMNH.RENA 6011 1Testudo Vosmaeri Suckow 1798 Cylindraspis vosmaeri RMNH.RENA 6001 1Trionychidae Trionyx stellatus var. japonicus Temminck & Schlegel 1834 Pelodiscus sinensis RMNH.RENA 3259, 3264 2 (species complex) Chelidae Batrachemys heliostemma McCord et al. 2001 Mesoclemmys heliostemma RMNH.RENA 31998-99 2Chelodina mccordi Rhodin 1994 Chelodina mccordi MCZ 176730 RMNH.RENA 10187; MCZ 17631-2 1Podocnemididae Emys Amazonica Spix 1824 Podocnemis expansa ZSM 2246/0/1 RMNH.RENA 3294; ZSM 7/0-14/0, 1

2446/0/2-7, 2447/0/1-4, 2730/0, 3095/0Emys macrocephala Spix 1824 Peltocephalus dumerilianus RMNH.RENA 6164 1CROCODYLIA Crocodylidae Crocodilus biporcatus raninus Müller & Schlegel 1841 Crocodylus raninus RMNH.RENA 3219 RMNH.RENA 37493 2Crocodilus biporcatus raninus Müller & Schlegel 1841 Crocodylus porosus RMNH.RENA 37489 1Crocodilus biporcatus raninus Müller & Schlegel 1841 Crocodylus siamensis RMNH.RENA 7939 + 21695; SMF 8090 1† Crocodilus ossifragus Dubois 1908 Crocodylus siamensis RMNH Dub. Skull and

many fragments manyCrocodilus (Gavialis) Schlegelii Müller 1838 Tomistoma schlegelii RMNH.RENA 3198, 3200,

7934-5, 35444-5, 35449, 39581 8Gavialidae † Garialus bengawanicus Dubois 1908 † Gavialus bengawanicus RMNH Dub. many fragments many


Table 1. Summary of nominal and present names, type status and number of type specimens of Testudines and Crocodylia in the RMNH collections.

Number in Original name Author Year Present name Holotype Lectotype Syntypes Para(lecto)types RMNH.RENATESTUDINES Chelydridae Chelonura Temminckii Troost in Harlan 1835 Macrochelys temminckii RMNH.RENA 6166 1Emys orbicularis occidentalis Fritz 1993 Emys orbicularis occidentalis RMNH.RENA 11371, 15003 3Geoemydidae Cyclemys enigmatica Fritz et al. 2008 Cyclemys enigmatica RMNH.RENA 3838, 6066, 4 6068, 27828 Cyclemys giebelii Hubrecht 1881 Notochelys platynota RMNH.RENA 3348 1Cyclemys pulchristriata Fritz et al. 1997 Cyclemys pulchristriata RMNH.RENA 4751 1Emys borneoensis Schlegel & Müller 1845 Batagur borneoensis RMNH.RENA 3296, 6210 2Emys dentata Gray 1831 Cyclemys dentata RMNH.RENA 6063, 6067, 40474 3Emys dentata Gray 1831 Cyclemys enigmatica RMNH.RENA 6062 1Emys subtrijuga Schlegel & Müller 1845 Malayemys subtrijuga RMNH.RENA 6082 RMNH.RENA 6084, 6085 3Emys vulgaris Gray 1831 Mauremys rivulata RMNH.RENA 3327 1Emys vulgaris japonica Temminck & Schlegel 1834 Mauremys japonica RMNH.RENA 3331-3334, 6

6142Emys vulgaris picta Schlegel 1844 Mauremys reevesii RMNH.RENA 3330 A-B, 2

MNHNP 1954 Geoemyda spengleri sinensis Fan 1931 Geoemyda spengleri RMNH.RENA 5889 1† Hardella isoclina Dubois 1908 † Mauremys? isoclina RMNH Dub. 2722 1Testudinidae Testudo dussumieri Gray 1831 Dipsochelys dussumieri RMNH.RENA 3231 1Testudo emys Schlegel & Müller 1840 Manouria emys emys RMNH.RENA 3808 RMNH.RENA 6005, 6030, 17967: 4 MNHNP 9422 Testudo Forstenii Schlegel & Müller 1845 Indotestudo forstenii RMNH.RENA 3811 1Testudo Strauchi Lidth de Jeude 1893 Psammobates geometricus RMNH.RENA 6011 1Testudo Vosmaeri Suckow 1798 Cylindraspis vosmaeri RMNH.RENA 6001 1Trionychidae Trionyx stellatus var. japonicus Temminck & Schlegel 1834 Pelodiscus sinensis RMNH.RENA 3259, 3264 2 (species complex) Chelidae Batrachemys heliostemma McCord et al. 2001 Mesoclemmys heliostemma RMNH.RENA 31998-99 2Chelodina mccordi Rhodin 1994 Chelodina mccordi MCZ 176730 RMNH.RENA 10187; MCZ 17631-2 1Podocnemididae Emys Amazonica Spix 1824 Podocnemis expansa ZSM 2246/0/1 RMNH.RENA 3294; ZSM 7/0-14/0, 1

2446/0/2-7, 2447/0/1-4, 2730/0, 3095/0Emys macrocephala Spix 1824 Peltocephalus dumerilianus RMNH.RENA 6164 1CROCODYLIA Crocodylidae Crocodilus biporcatus raninus Müller & Schlegel 1841 Crocodylus raninus RMNH.RENA 3219 RMNH.RENA 37493 2Crocodilus biporcatus raninus Müller & Schlegel 1841 Crocodylus porosus RMNH.RENA 37489 1Crocodilus biporcatus raninus Müller & Schlegel 1841 Crocodylus siamensis RMNH.RENA 7939 + 21695; SMF 8090 1† Crocodilus ossifragus Dubois 1908 Crocodylus siamensis RMNH Dub. Skull and

many fragments manyCrocodilus (Gavialis) Schlegelii Müller 1838 Tomistoma schlegelii RMNH.RENA 3198, 3200,

7934-5, 35444-5, 35449, 39581 8Gavialidae † Garialus bengawanicus Dubois 1908 † Gavialus bengawanicus RMNH Dub. many fragments many

Documents

Type specimens of recent and fossil Testudines and Crocodylia in the collections of the Netherlands Centre for Biodiversity Naturalis, Leiden, the Netherlands